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Premise During the last centuries, the area covered by urban landscapes is increasing all over the world. Urbanization can change local habitats and decrease connectivity among these habitats, with important consequences for species interactions. While several studies have found a major imprint of urbanization on plant–insect interactions, the effects of urbanization on seed predation remain largely unexplored. Methods We investigated the relative impact of sunlight exposure, leaf litter, and spatial connectivity on predation by moth and weevil larvae on acorns of the pedunculate oak across an urban landscape during 2018 and 2020. We also examined whether infestations by moths and weevils were independent of each other. Results While seed predation varied strongly among trees, seed predation was not related to differences in sunlight exposure, leaf litter, or spatial connectivity. Seed predation by moths and weevils was negatively correlated at the level of individual acorns in 2018, but positively correlated at the acorn and the tree level in 2020. Conclusions Our study sets the baseline expectation that urban seed predators are unaffected by differences in sunlight exposure, leaf litter, and spatial connectivity. Overall, our findings suggest that the impact of local and spatial factors on insects within an urban context may depend on the species guild. Understanding the impact of local and spatial factors on biodiversity, food web structure, and ecosystem functioning can provide valuable insights for urban planning and management strategies aimed at promoting urban insect diversity.

Premise: During the last centuries, the area covered by urban landscapes is increasing all over the world. Urbanization can change local habitats and decrease connectivity among these habitats, with important consequences for species interactions. While several studies have found a major imprint of urbanization on plant–insect interactions, the effects of urbanization on seed predation remain largely unexplored. Methods: We investigated the relative impact of sunlight exposure, leaf litter, and spatial connectivity on predation by moth and weevil larvae on acorns of the pedunculate oak across an urban landscape during 2018 and 2020. We also examined whether infestations by moths and weevils were independent of each other. Results: While seed predation varied strongly among trees, seed predation was not related to differences in sunlight exposure, leaf litter, or spatial connectivity. Seed predation by moths and weevils was negatively correlated at the level of individual acorns in 2018, but positively correlated at the acorn and the tree level in 2020. Conclusions: Our study sets the baseline expectation that urban seed predators are unaffected by differences in sunlight exposure, leaf litter, and spatial connectivity. Overall, our findings suggest that the impact of local and spatial factors on insects within an urban context may depend on the species guild. Understanding the impact of local and spatial factors on biodiversity, food web structure, and ecosystem functioning can provide valuable insights for urban planning and management strategies aimed at promoting urban insect diversity.

With climate change, plant-feeding insects increase their number of annual generations (voltinism). However, to what degree the emergence of a new herbivore generation affects the parasitism rate has not been explored. We performed a field experiment to test whether the parasitism rate differs between the first and the second generations of a specialist leaf miner (Tischeria ekebladella), both in the naturally univoltine and bivoltine parts of the leaf miner’s distribution. We found an interactive effect between herbivore generation and geographical range on the parasitism rate. The parasitism rate was higher in the first compared to the second host generation in the part of the range that is naturally univoltine, whereas it did not differ between generations in the bivoltine range. Our experiment highlights that shifts in herbivore voltinism might release top-down control, with potential consequences for natural and applied systems.

With climate change, plant-feeding insects increase their number of annual generations (voltinism). However, to what degree the emergence of a new herbivore generation affects the parasitism rate has not been explored. We performed a field experiment to test whether the parasitism rate differs between the first and the second generations of a specialist leaf miner (Tischeria ekebladella), both in the naturally univoltine and bivoltine parts of the leaf miner's distribution. We found an interactive effect between herbivore generation and geographical range on the parasitism rate. The parasitism rate was higher in the first compared to the second host generation in the part of the range that is naturally univoltine, whereas it did not differ between generations in the bivoltine range. Our experiment highlights that shifts in herbivore voltinism might release top-down control, with potential consequences for natural and applied systems.

Soil microbes play a key role in the regulation of plant performance. Rapid microbial responses to climate change (facilitated by their short life cycles) might help plant species (with lower short term adaptation capacity) to survive under increasing aridity. To test this hypothesis, we conducted a greenhouse experiment where seedlings of three tree species (Quercus suber, Quercus canariensis, Olea europaea) were grown with different soil inoculums and subjected to a process of drought and re-wetting. Soil inoculums were collected in a rainfall exclusion infrastructure located in Los Alcornocales Natural Park (Cádiz), where soils had been subjected to 30% rainfall reduction during four years. Half of the inoculums were sterilized to explore the net role of the soil microbiome in the seedlings response to drought. Results showed that four years of rainfall exclusion significantly altered the diversity and composition of the fungal and bacterial soil community. However, this alteration only translated into a higher ability to resist a subsequent drought in seedlings of O. europaea. Quercus seedlings were relatively independent of the soil microbiome in terms of growth, but not of physiological performance. Whereas Quercus seedlings growing in unsterilized soils were able to down-regulate photosynthesis and stomatal conductance in response to drought, seedlings in sterilized soils did not, which might compromise their long-term survival. Overall, our results showed that soil microbes play a relevant role in the response to drought of Mediterranean tree seedlings, and that they might contribute to the adaptation of at least some species to a drier climate.

1. Insect phenology consists of the timing of life events, as well as the number of generations (voltinism). While several studies have focused on the impact of climate on the timing of seasonal events, or the voltinism of single species, we have few insights into the factors that shape patterns of voltinism within ecological communities. Importantly, voltinism can have a major impact on population growth, species interactions, and rate of evolution. 2. We investigated the relative importance of spatial variation in temperature and species traits in shaping patterns of voltinism within an herbivore community feeding on deciduous oaks across a temperature gradient in Europe. 3. Voltinism increased with temperature, where the probability for a species to be univoltine decreased with temperature, whereas the probability for a species to be strictly multivoltine increased with temperature. The relative abundance of the first and subsequent generations of multivoltine species did not significantly change along the temperature gradient. Resource specialisation affected voltinism, where oligophagous and polyphagous species were more likely to be strictly multivoltine than narrow oligophagous species. Overwintering stage and body size did not affect voltinism, and there was no evidence that species traits influenced the relationship between temperature and voltinism. 4. Our findings highlight that temperature and species traits shape variation in voltinism within an herbivore community associated with oak trees. These temperature-induced shifts in voltinism within the oak-associated herbivore community may have profound effects on the synchrony within and between trophic levels, and consequently for food web structure and outbreak dynamics.

Plants interact with a large diversity of organisms, including insects and microorganisms. These species interactions are strongly influenced by climate, as illustrated by the advances in plant and insect phenology in response to increasing temperatures. Beyond changes in phenology, climate might also affect the number of times certain events take place during the same year, such as the number of leaf flushes of plants and the number of generations of insects (voltinism). In this thesis, I investigated the impact of the number of leaf flushes and insect voltinism on the oak food web. I focussed on the pedunculate oak Quercus robur and the community of herbivores and fungi that share oaks as a common resource. In the first half of my thesis, I looked into the impact of the co-existence of multiple leaf flushes on plant chemistry, insect attack, pathogen infection and the structure of the foliar fungal community along the distributional range of oaks in Europe. In the second half of my thesis, I examined the impact of temperature and species traits on the voltinism of the oak-associated herbivore community, as well as the joint impact of plant spring phenology and pathogen infection on the preference and performance of multigenerational attackers during the growing season. Overall, my findings showed that I) plant chemistry, insect attack and pathogen infection differ between leaf flushes, II) the foliar fungal community is strongly affected by oak leaf flush and latitude, III) temperature and resource specialisation are the main drivers for changes in voltinism within the oak-associated community of herbivores, and IV) spring phenology and pathogen infection affect the preference and performance of multigenerational attackers. Taken together, my thesis contributes to our understanding of the effects of climate-induced changes in the number of leaf flushes and voltinism on the structure and dynamics of the oak food web.

Aim: Leaves support a large diversity of fungi, which are known to cause plant diseases, induce plant defences or influence leaf senescence and decomposition. To advance our understanding of how foliar fungal communities are structured and assembled, we assessed to what extent leaf flush and latitude can explain the within- and among-tree variation in foliar fungal communities. Location: A latitudinal gradient spanning c. 20 degrees in latitude in Europe. Taxa: The foliar fungal community associated with a foundation tree species, the pedunculate oak Quercus robur. Methods: We examined the main and interactive effects of leaf flush and latitude on the foliar fungal community by sampling 20 populations of the pedunculate oak Quercus robur across the tree's range. We used the ITS region as a target for characterization of fungal communities using DNA metabarcoding. Results: Species composition, but not species richness, differed between leaf flushes. Across the latitudinal gradient, species richness was highest in the central part of the oak's distributional range, and foliar fungal community composition shifted along the latitudinal gradient. Among fungal guilds, the relative abundance of plant pathogens and mycoparasites was lower on the first leaf flush, and the relative abundance of plant pathogens and saprotrophs decreased with latitude. Conclusions: Changes in community composition between leaf flushes and along the latitudinal gradient were mostly a result of species turnover. Overall, our findings demonstrate that leaf flush and latitude explain 5%–22% of the small- and large-scale spatial variation in the foliar fungal community on a foundation tree within the temperate region. Using space-for-time substitution, we expect that foliar fungal community structure will change with climate warming, with an increase in the abundance of plant pathogens and mycoparasites at higher latitudes, with major consequences for plant health, species interactions and ecosystem

1. Climate change has been shown to advance spring phenology, increase the number of insect generations per year (multivoltinism) and increase pathogen infection levels. 2. However, we lack insights into the effects of plant spring phenology and the biotic environment on the preference and performance of multivoltine herbivores and whether such effects extend into the later part of the growing season. 3. To this aim, we used a multifactorial growth chamber experiment to examine the influence of spring phenology on plant pathogen infection, and how the independent and interactive effects of spring phenology and plant pathogen infection affect the preference and performance of multigenerational attackers (the leaf miner Tischeria ekebladella and the aphid Tuberculatus annulatus) on the pedunculate oak in the early, mid and late parts of the plant growing season. 4. Pathogen infection was highest on late phenology plants, irrespective of whether inoculations were conducted in the early, mid or late season. The leaf miner consistently preferred to oviposit on middle and late phenology plants, as well as healthy plants, during all parts of the growing season, whereas we detected an interactive effect between spring phenology and pathogen infection on the performance of the leaf miner. Aphids preferred healthy, late phenology plants during the early season, healthy plants during the mid season, and middle phenology plants during the late season, whereas aphid performance was consistently higher on healthy plants during all parts of the growing season. 5. Our findings highlight that the impact of spring phenology on pathogen infection and the preference and performance of insect herbivores is not restricted to the early season, but that its imprint is still present - and sometimes equally strong - during the peak and end of the growing season. Plant pathogens generally negatively affected herbivore preference and performance, and modulated the effects of spring phenology.

Many plant species produce multiple leaf flushes during the growing season, which might have major consequences for within-plant variation in chemistry and species interactions. Yet, we lack a theoretical or empirical framework for how differences among leaf flushes might shape variation in damage by insects and diseases. We assessed the impact of leaf flush identity on leaf chemistry, insect attack and pathogen infection on the pedunculate oak Quercus robur by sampling leaves from each leaf flush in 20 populations across seven European countries during an entire growing season. The first leaf flush had higher levels of primary compounds, and lower levels of secondary compounds, than the second flush, whereas plant chemistry was highly variable in the third flush. Insect attack decreased from the first to the third flush, whereas infection by oak powdery mildew was lowest on leaves from the first flush. The relationship between plant chemistry, insect attack and pathogen infection varied strongly among leaf flushes and seasons. Our findings demonstrate the importance of considering differences among leaf flushes for our understanding of within-tree variation in chemistry, insect attack and disease levels, something particularly relevant given the expected increase in the number of leaf flushes with climate change.

Many plant species produce multiple leaf flushes during the growing season, which might have major consequences for within-plant variation in chemistry and species interactions. Yet, we lack a theoretical or empirical framework for how differences among leaf flushes might shape variation in damage by insects and diseases. We assessed the impact of leaf flush identity on leaf chemistry, insect attack and pathogen infection on the pedunculate oak Quercus robur by sampling leaves from each leaf flush in 20 populations across seven European countries during an entire growing season. The first leaf flush had higher levels of primary compounds, and lower levels of secondary compounds, than the second flush, whereas plant chemistry was highly variable in the third flush. Insect attack decreased from the first to the third flush, whereas infection by oak powdery mildew was lowest on leaves from the first flush. The relationship between plant chemistry, insect attack and pathogen infection varied strongly among leaf flushes and seasons. Our findings demonstrate the importance of considering differences among leaf flushes for our understanding of within-tree variation in chemistry, insect attack and disease levels, something particularly relevant given the expected increase in the number of leaf flushes with climate change.

Aim Leaves support a large diversity of fungi, which are known to cause plant diseases, induce plant defences or influence leaf senescence and decomposition. To advance our understanding of how foliar fungal communities are structured and assembled, we assessed to what extent leaf flush and latitude can explain the within- and among-tree variation in foliar fungal communities. Location A latitudinal gradient spanning c. 20 degrees in latitude in Europe. Taxa The foliar fungal community associated with a foundation tree species, the pedunculate oak Quercus robur. Methods We examined the main and interactive effects of leaf flush and latitude on the foliar fungal community by sampling 20 populations of the pedunculate oak Quercus robur across the tree's range. We used the ITS region as a target for characterization of fungal communities using DNA metabarcoding. Results Species composition, but not species richness, differed between leaf flushes. Across the latitudinal gradient, species richness was highest in the central part of the oak's distributional range, and foliar fungal community composition shifted along the latitudinal gradient. Among fungal guilds, the relative abundance of plant pathogens and mycoparasites was lower on the first leaf flush, and the relative abundance of plant pathogens and saprotrophs decreased with latitude. Conclusions Changes in community composition between leaf flushes and along the latitudinal gradient were mostly a result of species turnover. Overall, our findings demonstrate that leaf flush and latitude explain 5%–22% of the small- and large-scale spatial variation in the foliar fungal community on a foundation tree within the temperate region. Using space-for-time substitution, we expect that foliar fungal community structure will change with climate warming, with an increase in the abundance of plant pathogens and mycoparasites at higher latitudes, with major consequences for plant health, species interactions and ecosystem dynam

The effects of climate change on oaks Quercus spp. constitute a main environmental concern for the conservation of temperate forests. In this context, we assessed the consequences of longer droughts on the interactions between the holm oak Quercus ilex L. and its main acorn pests. Infested acorns were prematurely abscised before reaching their potential size. The volume of the acorns attacked by Cydia fagiglandana (Lepidoptera) was smaller than those attacked by Curculio elephas (Coleoptera); however, their weight did not differ because Curculio larvae consumed more cotyledon. For the same reason, embryo survival likelihood was not lower in Cydia acorns despite their smaller size. Delays of late summer rain reduced infestation by Curculio, as soil hardness hampers adult emergence from their underground cells. By contrast, late and scarce precipitations benefited Cydia; rainfall might hamper adult flight and eggs/L1 larvae survival. There was not a “zero-sum” effect, because the decrease of Curculio infestation rates was not fully compensated by an increase of Cydia. Under the longer droughts projected for the Mediterranean Basin, our results predict lower infestation rates and higher acorn survival likelihood. However, further studies including other environmental factors are needed to better forecast the net consequences for holm oak fitness.

Premise Abiotic factors and plant species traits have been shown to drive latitudinal gradients in herbivory, and yet, population‐level factors have been largely overlooked within this context. One such factor is plant density, which may influence the strength of herbivory and may vary with latitude. Methods We measured insect herbivory and conspecific plant density (CPD) of oak (Quercus robur) seedlings and saplings along a 17° latitudinal gradient (2700 km) to test whether herbivory exhibited a latitudinal gradient, whether herbivory was associated with CPD, and whether such an association changed with latitude. Results We found a positive but saturating association between latitude and leaf herbivory. Furthermore, we found no significant relationship between CPD and herbivory, and such lack of density effects remained consistent throughout the sampled latitudinal gradient. Conclusions Despite the apparently negligible influence of plant density on herbivory for Q. robur, further research with other plant taxa and in different types of plant communities are needed to investigate density‐dependent processes shaping geographical variation in plant–herbivore interactions.

DNA barcoding identification needs a good characterization of intraspecific genetic divergence to establish the limits between species. Yet, the number of barcodes per species is many times low and geographically restricted. A poor coverage of the species distribution range may hamper identification, especially when undersampled areas host genetically distinct lineages. If so, the genetic distance between some query sequences and reference barcodes may exceed the maximum intraspecific threshold for unequivocal species assignation. Taking a group ofQuercusherbivores (moths) in Europe as model system, we found that the number of DNA barcodes from southern Europe is proportionally very low in the Barcoding of Life Data Systems. This geographical bias complicates the identification of southern query sequences, due to their high intraspecific genetic distance with respect to barcodes from higher latitudes. Pairwise intraspecific genetic divergence increased along with spatial distance, but was higher when at least one of the sampling sites was in southern Europe. Accordingly, GMYC (General Mixed Yule Coalescent) single-threshold model retrieved clusters constituted exclusively by Iberian haplotypes, some of which could correspond to cryptic species. The number of putative species retrieved was more reliable than that of multiple-threshold GMYC but very similar to results from ABGD and jMOTU. Our results support GMYC as a key resource for species delimitation within poorly inventoried biogeographic regions in Europe, where historical factors (e.g., glaciations) have promoted genetic diversity and singularity. Future European DNA barcoding initiatives should be preferentially performed along latitudinal gradients, with special focus on southern peninsulas.

Hoverflies are frequently used as biodiversity indicators and are targets of ecological studies across Europe. How hoverfly diversity responds to ecological variables is essential for species and habitat conservation. The present study is a first attempt to assess the hoverfly diversity of deciduous woodlands of Pyrenean oak (Quercus pyrenaica) in La Vera (Central-Western Spain), a habitat in need of conservation and a region that is poorly known in terms of hoverflies. Hoverflies were sampled with hand net in a mature-woodland site, a young-woodland site, and two close-by grassland sites. A higher number of species and individuals was recorded in woodland sites than in grassland sites, with only two species exclusive to the grasslands and thus, suggesting that woodland sites might be acting as a hoverfly reservoir and source. The proportion of species with phytophagous and saprophagous larvae in woodland sites was higher than outside the woodlands, probably due to a higher availability of resources in woodland sites for phytophagous and saprophagous species. The mature woodland was found to have a higher number of species and individuals than the young woodland, possibly due to the presence of grassy clearings and rotting materials in the trees of the mature woodland. The highest number of species was found in the predatory guild (Syrphinae), and smaller-bodied species within this guild were found to be more frequent in grassland than in woodland sites; possible explanations for this result are discussed. Further investigation is required to confirm and expand the results and conclusions found in this initial study on the hoverfly community of the poorly studied Q. pyrenaica woodlands of Central-Western Spain.

BACKGROUND AND AIMS: Classic theory on geographical gradients in plant-herbivore interactions assumes that herbivore pressure and plant defences increase towards warmer and more stable climates found at lower latitudes. However, the generality of these expectations has been recently called into question by conflicting empirical evidence. One possible explanation for this ambiguity is that most studies have reported on patterns of either herbivory or plant defences whereas few have measured both, thus preventing a full understanding of the implications of observed patterns for plant-herbivore interactions. In addition, studies have typically not measured climatic factors affecting plant-herbivore interactions, despite their expected influence on plant and herbivore traits. METHODS: Here we tested for latitudinal variation in insect seed predation and seed traits putatively associated with insect attack across 36 Quercus robur populations distributed along a 20° latitudinal gradient. We then further investigated the associations between climatic factors, seed traits and seed predation to test for climate-based mechanisms of latitudinal variation in seed predation. KEY RESULTS: We found strong but contrasting latitudinal clines in seed predation and seed traits, whereby seed predation increased whereas seed phenolics and phosphorus decreased towards lower latitudes. We also found a strong direct association between temperature and seed predation, with the latter increasing towards warmer climates. In addition, temperature was negatively associated with seed traits, with populations at warmer sites having lower levels of total phenolics and phosphorus. In turn, these negative associations between temperature and seed traits led to a positive indirect association between temperature and seed predation. CONCLUSIONS: These results help unravel how plant-herbivore interactions play out along latitudinal gradients and expose the role of climate in driving these outcomes t

DNA barcoding identification needs a good characterization of intraspecific genetic divergence to establish the limits between species. Yet, the number of barcodes per species is many times low and geographically restricted. A poor coverage of the species distribution range may hamper identification, especially when undersampled areas host genetically distinct lineages. If so, the genetic distance between some query sequences and reference barcodes may exceed the maximum intraspecific threshold for unequivocal species assignation. Taking a group of Quercus herbivores (moths) in Europe as model system, we found that the number of DNA barcodes from southern Europe is proportionally very low in the Barcoding of Life Data Systems. This geographical bias complicates the identification of southern query sequences, due to their high intraspecific genetic distance with respect to barcodes from higher latitudes. Pairwise intraspecific genetic divergence increased along with spatial distance, but was higher when at least one of the sampling sites was in southern Europe. Accordingly, GMYC (General Mixed Yule Coalescent) single‐threshold model retrieved clusters constituted exclusively by Iberian haplotypes, some of which could correspond to cryptic species. The number of putative species retrieved was more reliable than that of multiple‐threshold GMYC but very similar to results from ABGD and jMOTU. Our results support GMYC as a key resource for species delimitation within poorly inventoried biogeographic regions in Europe, where historical factors (e.g., glaciations) have promoted genetic diversity and singularity. Future European DNA barcoding initiatives should be preferentially performed along latitudinal gradients, with special focus on southern peninsulas.

Hoverflies are frequently used as biodiversity indicators and are targets of ecological studies across Europe. How hoverfly diversity responds to ecological variables is essential for species and habitat conservation. The present study is a first attempt to assess the hoverfly diversity of deciduous woodlands of Pyrenean oak (Quercus pyrenaica) in La Vera (Central-Western Spain), a habitat in need of conservation and a region that is poorly known in terms of hoverflies. Hoverflies were sampled with hand net in a mature-woodland site, a young-woodland site, and two close-by grassland sites. A higher number of species and individuals was recorded in woodland sites than in grassland sites, with only two species exclusive to the grasslands and thus, suggesting that woodland sites might be acting as a hoverfly reservoir and source. The proportion of species with phytophagous and saprophagous larvae in woodland sites was higher than outside the woodlands, probably due to a higher availability of resources in woodland sites for phytophagous and saprophagous species. The mature woodland was found to have a higher number of species and individuals than the young woodland, possibly due to the presence of grassy clearings and rotting materials in the trees of the mature woodland. The highest number of species was found in the predatory guild (Syrphinae), and smaller-bodied species within this guild were found to be more frequent in grassland than in woodland sites; possible explanations for this result are discussed. Further investigation is required to confirm and expand the results and conclusions found in this initial study on the hoverfly community of the poorly studied Q. pyrenaica woodlands of Central-Western Spain.