Joan Casanelles Abella

Land-use changes, driven by agricultural intensification and urbanization, are major contributors to biodiversity loss, altering habitats and reducing available resources. These changes impact species’ foraging strategies, particularly in human-modified ecosystems. While dietary shifts due to land-use changes have been well-studied in vertebrates, similar research in invertebrates, such as wild bees, remains limited. The present data paper provides a comprehensive dataset on the pollen collected from urban and rural populations of two bumblebee species (Bombus lapidarius and B. pascuorum) in Switzerland, examining pollen composition, nutrient content, and diet breadth. Additionally, by analyzing pollen from both body and leg-baskets, the dataset also offers a comprehensive overview of plant-bumblebee interactions. The data help understand plant-bumblebee interactions, pollination services, nutritional supply to larvae, and the impact of land-use changes on these processes. Furthermore, the dataset can be integrated with existing plant trait data to explore the effects of non-native species and other ecological factors on bumblebee foraging and nutrition in anthropogenically modified landscapes.

Cities can work towards addressing biodiversity conservation, climate adaptation, and social equity goals. Evidence indicates that cities can foster species conservation and human-nature interactions. Yet achieving multifunctional and equitable urban ecosystems requires addressing ecological gaps, enhancing monitoring methods, and engaging communities in sustainable urban planning. The Special Issue "Ecology for future cities" presented in this editorial, aims to contribute to solving existing gaps by investigating key themes shaping the field of urban ecology and emphasizing advancements to understand the social-ecological dynamics in cities. In this editorial paper, we present the seven papers of the special issue, and how they fall within six topics: (1) mapping urban ecosystems; (2) drivers and implications of human-nature interactions; (3) taxonomic coverage and biodiversity structure; (4) ecological transformations; (5) improving the temporal coverage; and (6) how ecology informs urban planning. We also provide future research directions based on these topics. We conclude the editorial with some personal thoughts regarding the two cities where we grew up, which have shaped our relationships to and understanding of urban ecosystems and contribute to motivating our research interests.

1. Urban ecosystems are formed by pronounced socio-ecological gradients, which are distinct from other ecosystems and can simultaneously filter and promote taxa, ultimately affecting their interactions. However, the strength of the effect of filtering and facilitation across the different trophic levels could vary among biotic and abiotic factors.
2. Here, we investigate the effects of habitat amount, temperature and host-enemy biotic interactions in shaping communities of cavity-nesting bees and wasps and their natural enemies. We installed trap-nests in 80 sites distributed along urban intensity gradients in five European cities (Antwerp, Paris, Poznan, Tartu and Zurich). We quantified the species richness and abundance of hosts and their natural enemies, as well as two performance traits (survival and parasitism) and two lifehistory traits (sex ratio and number of offspring per nest for the hosts). We analysed the importance of the abiotic and biotic variables using generalised linear models and multi-model inference.
3. We found that habitat amount was the main driver of multiple host responses, with larger habitat amounts resulting in higher species richness and abundance for hosts and natural enemies, as well as a larger probability of survival and a larger number of brood cells for hosts.
4. Local resources proxies shaped both bees and wasps and indicate different uses of existing vegetation between bees and wasps. Temperature proxies had a minor role in shaping host and natural enemies.
5. Biotic interactions were a main driver of host and enemy community structure, with natural enemies being strongly affected by host availability, that is, with direct density-dependence between hosts and their natural enemies. 6. Overall, our study highlights the importance of habitat amount and temperature in shaping urban food webs, as well as on biotic interactions through direct effects on hosts responses and the subsequent consequences for their natural enemies. As cities prepare to tackle the consequences of global change, strategies that make it possible to maintain habitat and mitigate urban overheating emerge as a key urban adaptation for biodiversity conservation.

In cities, green areas are essential for biodiversity conservation, with land cover heterogeneity being a decisive factor. Yet, as heterogeneity increases for a given green area, the patch size of land covers automatically decreases, as the area available for individual species, especially habitat specialist species. This relationship, known as the area-heterogeneity trade-off, is expected to lead to a unimodal relationship between species richness and land cover heterogeneity, and contrasted effects between species according to their level of urban avoidance.
We investigated the potential consequences of this trade-off on birds in green areas selected along an urban intensity gradient in six European cities. Using a European database on bird occurrences in nesting habitats, we defined a continuous gradient of urban avian avoidance.
We confirmed the marked area-heterogeneity trade-off in urban green areas but found no effect of land cover heterogeneity on total avian richness at green area level. However, both land cover heterogeneity and patch size were positively associated with richness of urban avoider species, indicating that urban avoiders fared better in green areas with large and heterogeneous patches. Total richness was also higher in green areas surrounded by an urban matrix composed of a variety of land covers.
To protect urban bird avoiders, which are most at risk in cities, green area managers and urban planners should thus be aware that land cover heterogeneity is not a panacea if patch sizes are too small. To conserve avian richness, we stress the importance of maintaining large vegetated areas as well as heterogeneity in land covers within the urban matrix.

Promoting urban green spaces is an effective strategy to increase biodiversity in cities. However, our understanding of how local and landscape factors influence trophic interactions in these urban contexts remains limited. Here, we sampled cavity-nesting bees and wasps and their natural enemies within 85 urban gardens in Zurich (Switzerland) to identify factors associated with the diversity and dissimilarity of antagonistic interactions in these communities. The proportions of built-up area and urban green area at small landscape scales (50 m radius), as well as the management intensity, sun exposure, plant richness and proportion of agricultural land at the landscape scale (250 m radius), were key drivers of interaction diversity. This increased interaction diversity resulted not only from the higher richness of host and natural enemy species, but also from species participating in more interactions. Furthermore, dissimilarity in community structure and interactions across gardens (beta-diversity) were primarily influenced by differences in built-up areas and urban green areas at the landscape scale, as well as by management intensity. Our study offers crucial insights for urban planning and conservation strategies, supporting sustainability goals by helping to understand the factors that shape insect communities and their trophic interactions in urban gardens.

Urban gardens have the potential to contribute to urban bee biodiversity. However, studies considering the effects of habitat amount and floral resources on bees using a functional approach, are largely lacking in Latin America, in particular in urban environments, where the potential competitive interactions between honeybees and wild bees deserve further attention. We evaluated how bee abundance and diversity, both taxonomic (richness, evenness) and functional (richness, evenness, and divergence of traits), are related to habitat amount in urban gardens (patch size, green cover in the surroundings), and the abundance and diversity of flowers (taxonomic and functional). We also examined the effect of the abundance of honeybees on urban wild bee communities. We selected 13 gardens within Córdoba city (Argentina) along an urbanization gradient based on vegetation cover, where we surveyed bee-flower interactions. We selected flower (i.e. morphology, color, and phenology) and bee (i.e. morphology, sociality, and diet) traits considered essential to plant-pollinator interactions to estimate flower and bee functional diversity, respectively. Bee taxonomic and functional diversity in urban gardens strongly depended on the floral species evenness, whereas the functional floral diversity and patch size, as well as the habitat amount at the landscape scale, did not significantly affect bee diversity. Moreover, the abundance of honeybees, once controlled by floral species evenness, did not influence the taxonomic structure of wild bee communities. Our results highlight that urban gardens can support functionally diverse bee communities, especially those with evenly distributed flower species.

Cities are socioecological systems that filter and select species, therefore establishing unique species assemblages and biotic interactions. Urban ecosystems can host richer wild bee communities than highly intensified agricultural areas, specifically in resource-rich urban green spaces such as allotments and family gardens. At the same time, urban beekeeping has boomed in many European cities, raising concerns that the fast addition of a large number of managed bees could deplete the existing floral resources, triggering competition between wild bees and honeybees. Here, we studied the interplay between resource availability and the number of honeybees at local and landscape scales and how this relationship influences wild bee diversity. We collected wild bees and honeybees in a pollination experiment using four standardized plant species with distinct floral morphologies. We performed the experiment in 23 urban gardens in the city of Zurich (Switzerland), distributed along gradients of urban and local management intensity, and measured functional traits related to resource use. At each site, we quantified the feeding niche partitioning (calculated as the average distance in the multidimensional trait space) between the wild bee community and the honeybee population. Using multilevel structural equation models (SEM), we tested direct and indirect effects of resource availability, urban beekeeping, and wild bees on the community feeding niche partitioning. We found an increase in feeding niche partitioning with increasing wild bee species richness. Moreover, feeding niche partitioning tended to increase in experimental sites with lower resource availability at the landscape scale, which had lower abundances of honeybees. However, beekeeping intensity at the local and landscape scales did not directly influence community feeding niche partitioning or wild bee species richness. In addition, wild bee species richness was positively influenced by local resource availability, whereas local honeybee abundance was positively affected by landscape resource availability. Overall, these results suggest that direct competition for resources was not a main driver of the wild bee community. Due to the key role of resource availability in maintaining a diverse bee community, our study encourages cities to monitor floral resources to better manage urban beekeeping and help support urban pollinators.

Wild bees are critical for multiple ecosystem functions but are currently threatened. Understanding the determinants of the spatial distribution of wild bee diversity is a major research gap for their conservation. We modeled wild bee α and β taxonomic and functional diversity in Switzerland to uncover countrywide diversity patterns and determine the extent to which they provide complementary information, assess the importance of the different drivers structuring wild bee diversity, identify hotspots of wild bee diversity, and determine the overlap between diversity hotspots and the network of protected areas. We used site-level occurrence and trait data from 547 wild bee species across 3343 plots and calculated community attributes, including taxonomic diversity metrics, community mean trait values, and functional diversity metrics. We modeled their distribution with predictors describing gradients of climate, resource availability (vegetation), and anthropogenic influence (i.e., land-use types and beekeeping intensity). Wild bee diversity changed along gradients of climate and resource availability; high-elevation areas had lower functional and taxonomic α diversity, and xeric areas harbored more diverse bee communities. Functional and taxonomic β diversities diverged from this pattern, with high elevations hosting unique species and trait combinations. The proportion of diversity hotspots included in protected areas depended on the biodiversity facet, but most diversity hotspots occurred in unprotected land. Climate and resource availability gradients drove spatial patterns of wild bee diversity, resulting in lower overall diversity at higher elevations, but simultaneously greater taxonomic and functional uniqueness. This spatial mismatch among distinct biodiversity facets and the degree of overlap with protected areas is a challenge to wild bee conservation, especially in the face of global change, and calls for better integrating unprotected land. The application of spatial predictive models represents a valuable tool to aid the future development of protected areas and achieve wild bee conservation goals.

Biomagnetic monitoring increasingly is applied to assess particulate matter (PM) concentrations, mainly using plant leaves sampled in small geographical area and from a limited number of species. Here, the potential of magnetic analysis of urban tree trunk bark to discriminate between PM exposure levels was evaluated and bark magnetic variation was investigated at different spatial scales. Trunk bark was sampled from 684 urban trees of 39 genera in 173 urban green areas across six European cities. Samples were analysed magnetically for the Saturation isothermal remanent magnetisation (SIRM). The bark SIRM reflected well the PM exposure level at city and local scale, as the bark SIRM (i) differed between the cities in accordance with the mean atmospheric PM concentrations and (ii) increased with the cover of roads and industrial area around the trees. Furthermore, with increasing tree circumferences, the SIRM values increased, as a reflection of a tree age effect related to PM accumulation over time. Moreover, bark SIRM was higher at the side of the trunk facing the prevailing wind direction. Significant relationships between SIRM of different genera validate the possibility to combine bark SIRM from different genera to improve sampling resolution and coverage in biomagnetic studies. Thus, the SIRM signal of trunk bark from urban trees is a reliable proxy for atmospheric coarse to fine PM exposure in areas dominated by one PM source, as long as variation caused by genus, circumference and trunk side is taken into account.

Urban green spaces (UGSs) are important elements of urban landscapes. Woody vegetation is a key component of UGSs, providing many socio-ecological benefits such as habitat provision and human well-being. Knowing plant diversity and vegetation configuration that underpin urban ecosystem processes and functions is critical to maximize nature contributions to city dwellers. Here, we present a well-replicated multi-city study showing a detailed description of taxonomic and structural diversity of woody vegetation in 225 UGSs distributed across seven European cities along a NE-SW gradient. Our aim was to understand how UGSs attributes, including size and fragmentation, influence woody vegetation features. A total of 418 woody species belonging to 76 families were identified. UGS size displayed weak positive correlations with woody species richness, but a strong negative correlation with woody species density. Alien woody species were abundant in all cities (from 40% of all species recorded in Antwerp to 64% in Lisbon and Zurich). Among the native tree species we found a predominance of Pinus spp. in southern cities and Acer spp. in cooler climates. On average, tree canopies extent was 56% of UGSs. This paper provides insights on the plant diversity and woody vegetation composition in UGSs of different size, climate and urban planning history. Our results encourage and contribute to future urban ecology studies involving different taxa and ecosystem services as well as support effective urban planning and management practices.

Urban beekeeping is booming, heightening awareness of pollinator importance but also raising concerns that its fast growth might exceed existing resources and negatively impact urban biodiversity. To evaluate the magnitude of urban beekeeping growth and its sustainability, we analysed data on beehives and available resources in 14 Swiss cities in 2012-2018 and modelled the sustainability of urban beekeeping under different scenarios of available floral resources and existing carrying capacities. We found large increases in hives numbers across all cities from an average 6.48 hives per km² (3139 hives in total) in 2012 to an average 10.14 hives per km² (9370 in total) in 2018 and observed that available resources are insufficient to maintain present densities of beehives, which currently are unsustainable.

1. Urban ecosystems can sustain populations of wild bees, partly because of their rich native and exotic floral resources. A better understanding of the urban bee diet, particularly at the larval stage, is necessary to understand biotic interactions and feeding behaviour in urban ecosystems, and to promote bees by improving the management of urban floral resources.
2. We investigated the larval diet and distribution patterns of four solitary wild bee species with different diet specialization (i.e. Chelostoma florisomne, Osmia bicornis, Osmia cornuta and Hylaeus communis) along urban intensity gradients in five European cities (Antwerp, Paris, Poznan, Tartu and Zurich) using two complementary analyses. Specifically, using trap-nests and pollen metabarcoding techniques, we characterized the species’ larval diet, assessed diet consistency across cities, and modelled the distribution of wild bees using species distribution models (SDMs).
3. Our results demonstrate that urban wild bees display different successful strategies to exploit existing urban floral resources: not only broad generalism (i.e. H. communis), but also intermediate generalism, with some degree of diet conservatism at the plant family or genus level (i.e. O. cornuta and O. bicornis), or even strict specialization on widely available urban pollen hosts (i.e. C. florisomne). Furthermore, we detected important diet variation in H. communis, with a switch from a herbaceous pollen diet to a tree pollen diet with increasing urban intensity.
4. Species distribution modelling indicated that wild bee distribution ranges inside urban ecosystems ultimately depend on their degree of specialization, and that broader diets result in less sensitivity to urban intensity.
5. Policy implications. Satisfying larval dietary requirements is critical to preserving and enhancing wild bee distributions within urban gradients. For high to intermediate levels of feeding specialization, we found considerable consistency in the preferred plant families or genera across the studied cities, which could be generalized to other cities where these bees occur. Identifying larval floral preferences (e.g. using pollen metabarcoding) could be helpful for identifying key plant taxa and traits for bee survival and for improving strategies to develop bee-friendly cities.

Urbanization poses threats and opportunities for the biodiversity of wild bees. At the same time, cities can harbour diverse wild bee assemblages, partily due to the unique plant assemblages providing resources. While bee dietary preferences have been investigated in different studies, bee dietary studies have been done mostly in non urban ecosystems and data based on plant visitation observations or palynological techniques. This data set describes the larval food preferences four wild bee species (i.e. Chelostoma florisomne, Hylaeus communis, Osmica bicornis and Osmia cornuta) common in urban areas in five different European cities (i.e. Antwerp, Paris, Poznan, Tartu and Zurich). In addition, this data set also describes the larval food preferences from individuals of three wild bee genera (i.e. Chelostoma sp., Hylaeus sp, and Osmia sp.) that cloud not be identified to the species level. This data results from a European-level study aimed at understanding the effects of urbanization on biodiversity across different cities and cityscapes, and a Swiss project aimed at understanding the effects of urban ecosystems in wild bee feeding behaviour. Wild bees were sampled using standardized trap-nests in 80 sites (32 in Zurich and 12 in each of the remaining cities), selected following a double gradient of available habitat at local and landscape scales. Larval pollen was obtained from the bee nests and identified using DNA metabarcoding. The data provide the plant composition at species or genus level preferred by each bee. This unique data can be used for a wide array of research questions, including urban ecology (e.g. diversity of food sources along urban gradients), bee ecology (characterisation of bee feeding preferences) or comparative studies on urban evolution of behavioural traits between urban and non-urban sites. In addition, the data can be applied for informing urban planning and conservation strategies, particularly concerning the flower resources (e.g. importance of exotic species and thus management activities). This data set can be freely used for non-commercial purposes and this data paper should be cited if you use the data; we request collaboration with the data set contact person be considered if this data set represents an important part of the data analyzed in a study.

The phyllosphere harbours a diverse and specific bacterial community, which influences plant health and ecosystem functioning. In this study, we investigated the impact of urban green areas connectivity and size on the composition and diversity of phyllosphere bacterial communities. Hereto, we evaluated the diversity and composition of phyllosphere bacterial communities of 233 Platanus x acerifolia and Acer pseudoplatanus trees in 77 urban green areas throughout six European cities. The community composition and diversity significantly differed between cities but only to a limited extent between tree species. We could show that urban intensity correlated significantly with the community composition of phyllosphere bacteria. In particular, a significant correlation was found between the relative abundances for 29 out of the 50 most abundant families and the urban intensity: the abundances of classic phyllosphere families, such as Acetobacteraceae, Planctomycetes and Beijerinkiaceae, decreased with urban intensity (i.e. more abundant in areas with more green, lower air pollution and lower temperature), while those related to human activities, such as Enterobacteriaceae and Bacillaceae, increased with urban intensity. The results of this study suggest that phyllosphere bacterial communities in European cities are associated with urban intensity and that effect is mediated by several combined stress factors.

To create more resilient cities, it is important that we understand the effects of the global change drivers in cities. Biodiversity-based ecological indicators (EIs) can be used for this, as biodiversity is the basis of ecosystem structure, composition, and function. In previous studies, lichens have been used as EIs to monitor the effects of global change drivers in an urban context, but only in single-city studies. Thus, we currently do not understand how lichens are affected by drivers that work on a broader scale. Therefore, our aim was to quantify the variance in lichen biodiversity-based metrics (taxonomic and trait-based) that can be explained by environmental drivers working on a broad spatial scale, in an urban context where local drivers are superimposed. To this end, we performed an unprecedented effort to sample epiphytic lichens in 219 green spaces across a continental gradient from Portugal to Estonia. Twenty-six broad-scale drivers were retrieved, including air pollution and bio-climatic variables, and their dimensionality reduced by means of a principal component analysis (PCA). Thirty-eight lichen metrics were then modelled against the scores of the first two axes of each PCA, and their variance partitioned into pollution and climate components. For the first time, we determined that 15% of the metric variance was explained by broad-scale drivers, with broad-scale air pollution showing more importance than climate across the majority of metrics. Taxonomic metrics were better explained by air pollution, as expected, while climate did not surpass air pollution in any of the trait-based metric groups. Consequently, 85% of the metric variance was shown to occur at the local scale. This suggests that further work is necessary to decipher the effects of climate change. Furthermore, although drivers working within cities are prevailing, both spatial scales must be considered simultaneously if we are to use lichens as EIs in cities at continental to global scales.

This article summarizes the data of a survey of flowering plants in 80 sites in five European cities and urban agglomerations (Antwerp, Belgium; greater Paris, France; Poznan, Poland; Tartu, Estonia; and Zurich, Switzerland). Sampling sites were selected based on a double orthogonal gradient of size and connectivity and were urban green areas (e.g. parks, cemeteries). To characterize the flowering plants, two sampling methodologies were applied between April and July 2018. First, a floristic inventory of the occurrence of all flowering plants in the five cities. Second, flower counts in sampling plots of standardized size (1 m²) only in Zurich. We sampled 2146 plant species (contained in 824 genera and 137 families) and across the five cities. For each plant species, we provide its origin status (i.e. whether the plants are native from Europe or not) and 11 functional traits potentially important for plant-pollinator interactions. For each study site, we provide the number of species, genera, and families recorded, the Shannon diversity as well as the proportion of exotic species, herbs, shrubs and trees. In addition, we provide information on the patch size, connectivity, and urban intensity, using four remote sensing-based proxies measured at 100- and 800-m radii.

Cities are human dominated ecosystems providing novel conditions for organisms. Research on urban biodiversity is rapidly increasing, yet it is still hampered by the partial spatial coverage of cities and because of existing taxonomic biases. Predictive models have proved to be a key tool to solve this shortfall. However, predictive models have rarely been used in urban ecosystems due to either the lack of sufficient species records or high-quality predictors (e.g. meaningful ecological maps). Here, we assemble a large cross-taxa inventory of 1446 species from 12 taxonomic groups, including several understudied invertebrate groups, sampled in 251 sites in Zürich, Switzerland. We investigate the species diversity distributions and the structure of species assemblages along artificial urban ecological gradients by applying predictive models. We find that the general species diversity distribution law, where assemblages are dominated by a few very abundant and frequent species, applied consistently across all taxonomic groups (3% of the species accounting for approximately 50% of abundance). Furthermore, only species of intermediate abundance and frequency are spatially structured along urban intensity gradients, with rare species numbers keeping constant even in the most urbanised parts of the city. In addition, we show that green areas with low mowing regimes are associated with higher species diversity in the majority of taxonomic groups. Hence, this suggests management relaxation as a low-cost solution to promote species richness. Our study demonstrates the potential of predictive modelling for addressing ecological questions in urban environments and to inform management and planning.

Cities are challenging environments for human life, because of multiple environmental issues driven by urbanization. These can sometimes be mitigated through ecosystem services provided by different functions supported by biodiversity. However, biodiversity in cities is affected by numerous factors, namely habitat loss, degradation, and fragmentation, as well as pollution, altered climate, and new biotic challenges. To better understand the link between biodiversity and ecosystem functions and services, we need to improve our mechanistic knowledge of these relationships. Trait-based ecology is a promising approach for unravelling the causes and consequences of biodiversity filtering on ecosystem processes and underlying services, but large gaps remain unexplored.
Here, we present a series of research directions that are aimed at extending the current knowledge of the relationship between trait-based biodiversity and ecosystem functions and services in cities. These directions are based on: (1) improving urban habitat mapping; (2) considering often neglected urban habitats and ecological niches; (3) integrating multiple urban gradients; (4) using trait-based approaches to improve our mechanistic understanding of the relationships between biodiversity and ecosystem functions and services; and (5) extending the involvement of citizens.
Pursuing these research directions may support the sustainable management of urban ecosystems and the long-term provision of ecosystem services, ultimately enhancing the well-being of urban populations.