Aim: Climate is a major driver of large-scale variability in biodiversity, as a likely result of more intense biotic interactions under warmer conditions. This idea fuelled decades of research on plant-herbivore interactions, but much less is known about higher-level trophic interactions. We addressed this research gap by characterizing both bird diversity and avian predation along a climatic gradient at the European scale. Location: Europe. Taxon: Insectivorous birds and pedunculate oaks. Methods: We deployed plasticine caterpillars in 138 oak trees in 47 sites along a 19 degrees latitudinal gradient in Europe to quantify bird insectivory through predation attempts. In addition, we used passive acoustic monitoring to (i) characterize the acoustic diversity of surrounding soundscapes; (ii) approximate bird abundance and activity through passive acoustic recordings; and (iii) infer both taxonomic and functional diversity of insectivorous birds from recordings. Results: The functional diversity of insectivorous birds increased with warmer climates. Bird predation increased with forest cover and bird acoustic activity but decreased with mean annual temperature and functional richness of insectivorous birds. Contrary to our predictions, climatic clines in bird predation attempts were not directly mediated by changes in insectivorous bird diversity or acoustic activity, but climate and habitat still had independent effects on predation attempts. Main Conclusions: Our study supports the hypothesis of an increase in the diversity of insectivorous birds towards warmer climates but refutes the idea that an increase in diversity would lead to more predation and advocates for better accounting for activity and abundance of insectivorous birds when studying the large-scale variation in insect-tree interactions.

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. 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. 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.

The COVID-19 shutdown has caused a quasi-experimental situation for ecologists in Spring 2020, providing an unprecedented release in acoustic space for avian soundscapes due to the lowest technophony levels experienced for decades. We conducted large-scale passive acoustic monitoring in 68 forest stands during and after the shutdown to compare their acoustic diversity under different management regimes. We designed a before-after sampling scheme of 18 paired stands to evaluate the short-term effect of shutdown on diel and nocturnal acoustic diversity of forest soundscapes. We assessed whether old-growth preserves hosted higher acoustic diversity and vocal activity of flagship specialist birds than production stands during the shutdown, and whether the effect of management was mediated by landscape fragmentation and distance to roads. We derived acoustic richness and vocal activity of flagship specialist birds by systematically performing 15-min long aural listening to identify species vocalizations from all recorded stands. The end of the COVID-19 shutdown led to a rapid decrease in diel and nocturnal biophony and acoustic diversity. During the shutdown, we found significantly higher biophony and acoustic diversity in old-growth preserves than in production stands. Bird acoustic richness and vocalizations of the two most frequent flagship specialists, Dendrocoptes medius and Phylloscopus sibilatrix, were also both higher in old-growth stands. Interestingly, this positive effect of old-growth stands on forest soundscapes suggested that they could potentially attenuate traffic noise, because the distance to roads decreased acoustic diversity and biophony only outside old-growth preserves. Similarly, flagship bird richness increased with old-growth cover in the surrounding landscape while edge density had a negative effect on both acoustic diversity and flagship birds. We suggest that enhancing the old-growth preserve network implemented across French public forests would provide a connected frame of acoustic sanctuaries mitigating the ever-increasing effect of technophony on the acoustic diversity of temperate forest soundscapes.

Insect herbivores co-occurring on the same host plant interact in various ways. In particular, early-season insect herbivory triggers a wide range of plant responses that can determine the performance of herbivores colonizing the plant later in the course of the season. But the strength and direction of such effects are debated, and virtually unknown in the case of novel interactions involving exotic insects in their introduction range. We conducted an observational field study in SW France, a region recently invaded by the Oak Lace Bug (OLB, Corythucha arcuata Say). We measured early chewing damage and subsequent OLB damage in four oak species (Quercus robur, Q. pubescens, Q. cerris and Q. ilex). We set up a complementary non-choice experiment in the laboratory, feeding OLB with leaves with or without prior herbivory. The four oak species differed in their sensitivity to OLB damage, Q. ilex being broadly resistant. Prior herbivory promoted OLB damage in the laboratory experiment, but not in the field. However, prior herbivory did not alter the rank of oak resistance to the OLB. Our results suggest possible synergistic effects between spring defoliators and the OLB. This study brings insight into herbivore-herbivore interactions and their possible implications for forest management.

Experimental studies of biotic interactions in real field conditions are essential to understand the structure and functioning of ecological networks. The use of artificial caterpillars to mimic actual prey availability is generally seen as a standard approach to compare the activity and diversity of predators along environmental gradients. Yet, even with standardized material and procedures, biases may still affect data from multiple observers with different expertise. We used pictures of artificial caterpillars with or without various predation marks, in an online survey that was targeted for the participants of the project, to evaluate the reliability of predation marks identification made by non-scientists and by scientists with and without previous experience in predation mark identification. Pictures of artificial caterpillars displayed typical marks left by birds, mammals and arthropods, as well as non-predation marks (‘false positive’). 357 respondents scanned 7140 pictures of these pictures. Self-declared scientists were more confident and accurate in their observations than non-scientists, but the differences in correct identifications among scientists and non-scientists were low. Self-declared scientists with previous experience were also more accurate than scientists without previous experience, while there were no differences in self-confidence among scientists with and without previous experience. Accuracy in predation mark identification did not differ among types of predators, but respondents were more keen to identify marks left by birds or mammals than arthropods. Our results have practical implications for the design of multi-observer projects relying on artificial caterpillars as a proxy to assess predation intensity, in particular in the context of citizen science.

Urbanization is recognized as an important driver of the diversity and abundance of tree associated insect herbivores, but its consequences for insect herbivory are controversial. 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 effect of urbanization and local canopy cover on insect herbivory in the pedunculate oak (Quercus robur) throughout most of its geographic range in Europe. The damage caused by chewing insect herbivores as well as the incidence of leaf-mining and gall-inducing herbivores consistently decreased with increasing urbanization around focal oaks. Herbivory by chewing herbivores increased with increasing forest cover, regardless of urbanization. In contrast, an increase in local canopy cover buffered the negative effect of urbanization on leaf-miners and strengthened its effect on gall-inducers. These results show the complexity of plant-herbivore interactions in urbanized areas, highlighting that the presence of local canopy cover within cities has the potential to attenuate or modify the effect of urbanization on biotic interactions.

Aim The strength of species interactions is traditionally expected to become stronger toward the Equator. However, recent studies have reported opposite or inconsistent latitudinal trends in the bottom-up (plant quality) and top-down (natural enemies) forces driving insect herbivory, possibly because these forces have rarely been studied concomitantly. This makes previous attempts to understand the effect of large scale climatic gradients on insect herbivory unsuccessful. Location Europe Time period 2018-2019 Major taxa studied Quercus robur Methods We used scholar-based citizen science to simultaneously test for latitudinal variation in plant-herbivore-natural enemy interactions. We further investigated the underlying climatic factors associated with variation in herbivory, leaf chemistry and attack rates in Quercus robur across its complete latitudinal range in Europe. We quantified insect herbivory and the occurrence of specialist herbivores as well as leaf chemistry and bird attack rates on dummy caterpillars on 261 oak trees. Results Climatic factors rather than latitude per se were the best predictors of the large-scale (geographical) variation in the incidence of gall-inducers and leaf-miners as well as of leaf nutritional quality. However, insect herbivory, plant chemical defences (leaf phenolics) and bird attack rates were not influenced by latitude or climatic factors. The incidence of leaf-miners increased with increasing concentrations of hydrolysable tannins and decreased with those of condensed tannins, whereas the incidence of gall-inducers increased with increasing leaf soluble sugar concentration and decreased with increasing leaf C:N ratios. However, neither other traits nor bird attack rates varied with insect herbivory. Main conclusions These findings help to refine our understanding of the bottom-up and top-down mechanisms driving geographical variation in plant-herbivore interactions, and urge for further examination of the drivers of insect herbivory on trees.

The presence of heterospecific neighbours can affect damage caused by pest insects on focal plants. However, how plant neighbours influence herbivore performance is poorly understood. We tested the independent and interactive effects of tree species identity and tree neighbour type (conspecific vs. heterospecific) on the performance of a major oak pest, the oak processionary moth larvae (OPM, Thaumetopoea processionea ) fed on Quercus robur and Q. petraea . We performed a factorial greenhouse experiment in which we grew two oak saplings per pot, either from the same species or from both species. We quantified growth and mortality of OPM larvae, leaf phenolic compounds, C:N ratio and bud phenology. OPM larvae performed significantly better on Q. petraea than on Q. robur , regardless of plant neighbour type. Phenolic compounds and C:N, but not phenology, differed between oak species and neighbour species identity. Only bud phenology had a significant effect on OPM performance, which was better when young larvae had access to recently unfolded leaves, regardless of oak species and neighbour identity. Although oak neighbour identity altered the expression of leaf traits, this effect had no measurable consequences on OPM performance. However, further studies should consider the effect of oak species neighbour on OPM preferences for either Q. robur or Q. petraea , in pure and mixed stands, before translating current results into recommendations for forest management. Author Contribution TD and BC conceived the study and acquired the data. TD performed experiment and analysed the data. EV, TD and XM performed the chemical analyses. TD and BC drafted the first version of the manuscript and all authors wrote the final version of the manuscript. TD, BC conceived and designed research. TD performed experiment and analysed data. EV, TD, XM performed the chemical analyses. TD, BC wrote the manuscript. All authors read and approved the manuscript. AM and DB conceived and designed research. AM and BB conducted experiments. GR contributed new reagents or analytical tools. AM, BB and GR analysed data. AM wrote the manuscript. All authors read and approved the manuscript.

Abstract European forests are expanding and becoming denser following the widespread abandonment of farmland and rural areas. Spontaneous forest regrowth provides a cost‐effective opportunity to restore ecosystems, enhance multifunctionality and sustainability and mitigate climate change. Yet, little is known about the goods and services that such forests provide to people. We assessed the changes in nature's contributions to people (NCP) from spontaneous forest regrowth, i.e. forest expansion and densification, in South‐West Europe. We investigated 65 forest plots in four different landscapes with contrasting ecological and societal contexts. Two landscapes are located in rural areas undergoing human exodus and forest expansion and densification; the other two, in peri‐urban areas with intense land use and forest densification but negligible expansion. For each forest plot, we estimated variables related to ten out of the 18 main NCP defined by the Intergovernmental Science‐Policy Platform on Biodiversity and Ecosystem Services (IPBES). Regulating and material NCP were addressed using variables measured in the field as proxies. Non‐material NCP were studied through stakeholder interviews. Our results show across the cases that forest expansion and densification are generally associated with greater climate regulation and energy provision. Changes in other NCP, especially in non‐material ones, were strongly context‐dependent. The social perception of spontaneous forest regrowth was primarily negative in rural areas and more positive in peri‐urban landscapes. Passive restoration through spontaneous forest expansion and densification can enhance regulating and material NCP, especially when adaptive management is applied. To optimise NCP and to increase the societal awareness of and interest in spontaneous forest regrowth, the effects of this process should be analysed in close coordination with local stakeholders to unveil and quantify the many and complex trade‐offs involved in rural or peri‐urban social perceptions. A free Plain Language Summary can be found within the Supporting Information of this article.

Background and Aims Highly controlled experiments revealed that plant genetic diversity and relatedness can shape herbivore communities and patterns of herbivory. Evidence from the field is scarce and inconsistent. We assessed whether a genetic signal underlying herbivory can be detected in oak forest stands when accounting for variation at smaller (within-tree) and larger (among-stand) scales. Methods We tested relationships between tree genetic relatedness, leaf chemical defences and insect herbivory at different canopy layers in 240 trees from 15 Pedunculate oak ( Quercus robur ) forest stands and partitioned sources of variability in herbivory and defences among stands, individuals, and branches. Key Results Leaf defences, insect herbivory, and their relationship differed systematically between the upper and the lower tree canopy. When accounting for this canopy effect, the variation explained by tree genetic relatedness rose from 2.8 to 34.1 % for herbivory and from 7.1 to 13.8 % for leaf defences. The effect was driven by markedly stronger relationships in the upper canopy. Conclusions Our findings illustrate that properly accounting for other sources of variation acting at different scales can reveal potentially relevant effects of the host plant genotype on patterns of leaf chemical defences and associated insect herbivory in natural tree populations.

Key message Analyses of dendrochronological data from 15 recently established tablished stands of pedunculate oak (Quercus robur L.) revealed that functions describing potential tree growth in the absence of neighbours varied more between stands than functions describing competitive effects of conspecific neighbours. This suggests that competition functions can more easily be transferred among stands than potential growth functions. Context The variability inherent in the natural establishment of tree stands raises the question whether one can find general models for potential growth and competition that hold across stands.AimsWe investigated variation in potential growth and competition among recently established stands of Q. robur and tested whether this variation depends on stand structure. We also tested whether competition is symmetric or asymmetric and whether it is density-dependent or size-dependent. Lastly, we examined whether between-year growth variation is synchronous among stands. Methods Potential growth, competition and between-year growth variation were quantified with statistical neighbourhood models. Model parameters were estimated separately for each stand using exhaustive mapping and dendrochronology data. Results Competition was best described with an asymmetric size-dependent model. Functions describing potential growth varied more among forest stands than competition functions. Parameters determining these functions could not be explained by stand structure. Moreover, annual growth rates showed only moderate synchrony across stands. Conclusion The substantial between-stand variability in potential growth needs to be considered when assessing the functioning, ecosystem services and management of recently established Q. robur stands. In contrast, the relative constancy of competition functions should facilitate their extrapolation across stands.

During the past decades, a multitude of oak stands have spontaneously established across the pine-dominated landscapes of the French Landes de Gascogne. Yet their future performance under modern climate change is unknown. We show that coppiced, dominant trees are most prepared to cope with drought episodes, displaying higher basal area increment and lower sensitivity to extreme events. Forest stands dominated by pedunculate oak (Quercus robur L.) have spontaneously established across the pine-dominated landscapes of the French Landes de Gascogne. These oak stands are typically unmanaged and unsystematically coppiced, resulting in mixtures of single- and multi-stemmed (coppiced) trees. To determine the ability of spontaneous oak forest stands to face climate change–related hazards, by analysing differences in growth (tree-ring width and basal area increment—BAI), wood density and climate sensitivity depending on their tree architecture (single- vs multi-stemmed trees) and their social status in the forest. We exhaustively cored 15 oak stands (n = 657 trees). We compared stand characteristics and climate sensitivity between tree architectures considering two sampling designs, either all sampled trees (the exhaustive sampling) or those with a dominant status (dominant sampling). At the tree level, we used linear mixed effects models to compare wood density and growth between tree architectures and the trees’ social status within the canopy layer (dominant- vs non-dominant trees). Multi-stemmed trees exhibited higher wood density than single-stemmed trees for diameters > 30 cm. Dominant multi-stemmed trees showed lower sensitivity to extreme events (pointer years), higher BAI but lower annual growth rates than dominant single-stemmed trees. Dominant multi-stemmed trees are potentially the most prepared ones to cope with increasing soil water deficit following drought episodes, at least during the first 60 years of the life of the tree. The vulnerability to face harsher climate conditions for Q. robur stands can be misled when using a dominant sampling design.

Scientific knowledge in the field of ecology is increasingly enriched by data acquired by the general public participating in citizen science (CS) programs. Yet, doubts remain about the reliability of such data, in particular when acquired by schoolchildren. We built upon an ongoing CS program, 'Oak Bodyguards', to assess the ability of schoolchildren to accurately estimate the strength of biotic interactions in terrestrial ecosystems. We used standardized protocols to estimate attack rates on artificial caterpillars and insect herbivory on oak leaves. We compared estimates made by schoolchildren with estimates made by professional scientists who had been trained in predation and herbivory assessments (henceforth, trained scientists), and trained scientists’ estimates with those made by professional scientists with or without expertise (untrained) in predation or herbivory assessment. Compared with trained scientists, both schoolchildren and untrained professional scientists overestimated attack rates, but assessments made by the latter were more consistent. Schoolchildren tended to overestimate insect herbivory, as did untrained professional scientists. Raw data acquired by schoolchildren participating in CS programs therefore require several quality checks by trained professional scientists before being used. However, such data are of no less value than data collected by untrained professional scientists. CS with schoolchildren can be a valuable tool for carrying out ecological research, provided that the data itself is acquired by professional scientists from material collected by citizens.

Nutrient availability is important for plant community composition and diversity, but most studies focus on inorganic nutrients. Far less is known about the impact of nutrients in organic forms such as herbivore dung. Here we show that dung of 11 European herbivore species varies widely in nitrogen (N) and phosphorus (P) concentrations, as well as in C:N:P ratios. We demonstrate that variation in dung quality of five herbivore species influences the diversity and composition of a mesocosm plant community. The impact of dung quality was at least as strong as, or stronger than, the effect of manipulating the quantity of dung by a factor six. Our study supports the hypothesis that both nutrient quantity and nutrient imbalances are important controlling factors for plant species diversity, and stresses the important role of herbivores on plant communities, not only via selective foraging, but also via stoichiometric variation of nutrients in their dung.

The value of novel native broadleaf woodlands for biodiversity conservation is important to consider for adequate forest management in rural landscapes. Passive reforestation has been proposed as a cost-efficient tool for creating networks of novel native forest stands that would help restoring biodiversity and associated ecosystem services. Yet to date the ecological functioning of such stands remains strongly understudied compared to forest remnants resulting from longer-term fragmentation. We assessed how the size and connectivity of newly established Pedunculate oak (Quercus robur L.) stands in rural landscapes of SW France affect rates of herbivory by different insect guilds as well as rates of avian insectivory and the abundance and richness of insectivorous birds. Comparing 18 novel forest stands along a gradient of size (0.04-1.15 ha) and cover of broadleaf forests in the surroundings (0-30% within a 500 radius), we found that even the smallest stands are colonised by leaf miners and chewers/skeletonizers, and that rates of herbivory are globally comparable to those reported from older and larger oak forests. The size of stands had a relatively minor effect on herbivory, whereas it increased the abundance of insectivorous bird. It also determined rates of avian insectivory as estimated by an experiment with plasticine caterpillars. These rates were however rather low and unrelated with the extent of herbivory in the stand. Overall, our study indicates that insect herbivores tend to react more rapidly to the establishment of novel native forests than their avian predators as the latter may depend on the development of larger patches of suitable habitat in the surrounding landscape. To favour a rapid build-up of diverse, and hence stable, trophic networks involving insect herbivores and their predators, woodland creation schemes should therefore primarily focus on habitat size and quality.

The 'Oak bodyguards' citizen science project aimed to assess the effects of climate on two key biotic interactions occurring widely in natural and anthropized ecosystems, i.e., the top-down and bottom-up forces controlling insect herbivory on leaves of the English oak (Quercus robur) trees. We chose the English oak as a model species as it is one of the most common and most emblematic forest trees in Europe, with a geographic range spanning more than 19 degrees of latitude. It is also widespread in both rural, suburban and urban environments. We exposed dummy plasticine caterpillars in trees to estimate predation rates. We additionally estimated leaf insect herbivory as the percentage of leaf area removed or damaged by insect herbivores. The Oak bodyguards project involved professional scientists and school children throughout Europe. We designed a simple protocol that was applied by both partner types. The protocol was written by scientists in collaboration with science instructors and communication officers. It was available in five languages: French, English, German, Spanish and Portuguese. The present document is the English version of the protocol. It describes how to estimate insect herbivory on oak leaves as well as predation rate on artificial prey made with green modelling clay.

Tree diversity has long been recognized as a major driver of insect herbivory in forest ecosystems. However, predicting the strength and direction of tree diversity effects in real-world situations has proven elusive. One likely reason is that most studies have focused on within-stand dynamics and insufficiently captured other ecological drivers of insect herbivory that can act at broader (i.e., landscape) and finer (i.e., individual trees) scales. We measured herbivory as leaf area consumed by insect herbivores in pedunculate oaks (Quercus robur) growing in mixed and pure forest stands in southwestern France. We assessed the effects of oak spatial isolation within the landscape, tree stand diversity, forest canopy stratification as well as the influence of leaf traits on insect herbivory. Insect herbivory increased with stand isolation regardless of tree diversity. Diversity effects were contingent upon the canopy stratum as insect herbivory in mixed stands exceeded that of pure stands only in the upper stratum. Leaf traits varied between pure and mixed stands and among canopy strata. Insect herbivory was negatively correlated with LDMC and positively with SLA. However, the observed effects of tree diversity, canopy stratum and stand isolation on insect herbivory were only partially driven by variability in oak leaf traits. Our findings illustrate that, in real-world contexts, insect herbivory can be driven by a complex interplay of multiple, scale-dependent drivers. They help step forward towards a more profound understanding of the complex forces drive insect herbivory in managed forest ecosystems.

Woody plant encroachment has increased across the globe and threatens biodiversity associated with open habitats. In order to prevent or reduce woody encroachment, conservation managers across Europe introduce large mammalian herbivores. While up to recently, managers were mostly using free-ranging domestic cattle and horses for this, there is an increasing interest in the use of European bison for nature management. However, we lack studies that compare the impact of these different grazers on vegetation. We report results from a unique grazing pilot in the National Park Zuid-Kennemerland, a heterogeneous coastal dune landscape in the Netherlands, where European bison, horses, and cattle were introduced to reverse the encroachment of grass and shrub species. We present results of an 8-year study on the development of woody and grassy vegetation on fixed transects in three different grazing areas within the national park; one area with European bison and horses, one area with cattle and horses, and one area where these large grazers were excluded. In all three areas, rabbit, fallow deer, and roe deer were present. Over time, we observed strong reductions in the vitality of several woody species, such as spindle tree, and this decline was similar across all areas. Grass height and cover also declined and the proportion of herbs increased in all three grazing areas in similar ways. However, the type of herbivore use (debarking, foraging on buds, branches) of several woody species differed significantly among areas. For instance, maple tree was only debarked in the E. bison area, while hawthorn branches were eaten significantly more in the cattle than in the bison area. Due to differences in herbivore densities among areas, it was difficult to draw strong conclusions on how the different herbivore species differed in their impact, but, importantly, we found that grazing regimes with bison can lead to as strong effects on vegetation structure and composition as grazing regimes with cattle. This is an important result since certain conditions, such as legal aspects, may motivate managers to introduce a wild large grazer rather than a domesticated one.

Plants have compensating mechanisms to alleviate the loss of vegetative and reproductive tissues by herbivores. In this study, we evaluated how reproductive traits (size and production of fruit and seed, as well as germination and viability of seeds) of Myrtus communis (myrtle) were affected by goat grazing exclosure of forest understory in the Doñana Natural Park. One hundred and fifty myrtles were selected in three sites with similar environmental characteristics, although with a different length of grazing exclusion: (1) exclosure for 2 years (GE2); (2) exclosure for 3 years (GE3); and (3) long ago grazed plots (GE15): non-accessible to goat for at least 15 years. Fruit production, fruit and seed morphology and seed viability and germination were evaluated. Length of exclosure to goats increased fruit production, mass and volume of the fruits and seed, and endocarp mass. However, seed viability was higher in more recently grazed plots (70–90%) than in GE15 plots (51%). The final balance of viable/germinable seeds produced by an average sized myrtle in each treatment showed that the recently grazed myrtles (GE2) produced nearly five times less seeds (1200) than the myrtles grazed 15 years ago (5500). However, myrtles recovered the ability to produce viable seeds quickly, almost tripling, each year of non-grazing the number of viable/germinable seeds. The results of this study indicated that goat grazing could affect regeneration of myrtle because it reduced reproduction output and the number of viable propagules. Future studies should evaluate other related ecological aspects, such as fruit consumption by potential frugivores and seed dispersal (distance and place of seed deposition).