Body size is intrinsically linked to metabolic rate and life-history traits, and is a crucial determinant of food webs and community dynamics. The increased temperatures associated with the urban-heat-island effect result in increased metabolic costs and are expected to drive shifts to smaller body sizes . Urban environments are, however, also characterized by substantial habitat fragmentation , which favours mobile species. Here, using a replicated, spatially nested sampling design across ten animal taxonomic groups, we show that urban communities generally consist of smaller species. In addition, although we show urban warming for three habitat types and associated reduced community-weighted mean body sizes for four taxa, three taxa display a shift to larger species along the urbanization gradients. Our results show that the general trend towards smaller-sized species is overruled by filtering for larger species when there is positive covariation between size and dispersal, a process that can mitigate the low connectivity of ecological resources in urban settings . We thus demonstrate that the urban-heat-island effect and urban habitat fragmentation are associated with contrasting community-level shifts in body size that critically depend on the association between body size and dispersal. Because body size determines the structure and dynamics of ecological networks , such shifts may affect urban ecosystem function.

The increasing urbanization process is hypothesized to drastically alter (semi‐)natural environments with a concomitant major decline in species abundance and diversity. Yet, studies on this effect of urbanization, and the spatial scale at which it acts, are at present inconclusive due to the large heterogeneity in taxonomic groups and spatial scales at which this relationship has been investigated among studies. Comprehensive studies analysing this relationship across multiple animal groups and at multiple spatial scales are rare, hampering the assessment of how biodiversity generally responds to urbanization. We studied aquatic (cladocerans), limno‐terrestrial (bdelloid rotifers) and terrestrial (butterflies, ground beetles, ground‐ and web spiders, macro‐moths, orthopterans and snails) invertebrate groups using a hierarchical spatial design, wherein three local‐scale (200 m × 200 m) urbanization levels were repeatedly sampled across three landscape‐scale (3 km × 3 km) urbanization levels. We tested for local and landscape urbanization effects on abundance and species richness of each group, whereby total richness was partitioned into the average richness of local communities and the richness due to variation among local communities. Abundances of the terrestrial active dispersers declined in response to local urbanization, with reductions up to 85% for butterflies, while passive dispersers did not show any clear trend. Species richness also declined with increasing levels of urbanization, but responses were highly heterogeneous among the different groups with respect to the richness component and the spatial scale at which urbanization impacts richness. Depending on the group, species richness declined due to biotic homogenization and/or local species loss. This resulted in an overall decrease in total richness across groups in urban areas. These results provide strong support to the general negative impact of urbanization on abundance and species richness within habitat patches and highlight the importance of considering multiple spatial scales and taxa to assess the impacts of urbanization on biodiversity.

Traditionally metacommunity studies have quantified the relative importance of dispersal and environmental processes on observed β-diversity. Separating β-diversity into its replacement and nestedness components and linking such patterns to metacommunity drivers can provide richer insights into biodiversity organization across spatial scales. It is often very difficult to measure actual dispersal rates in the field and to define the boundaries of natural metacommunities. To overcome those limitations, we revisited an experimental metacommunity dataset to test the independent and interacting effects of environmental heterogeneity and dispersal on each component of β-diversity. We show that the balance between the replacement and nestedness components of β-diversity resulting from eutrophication changes completely depending on dispersal rates. Nutrient enrichment negatively affected local zooplankton diversity and generated a pattern of β-diversity derived from nestedness in unconnected, environmentally heterogeneous landscapes. Increasing dispersal erased the pattern of nestedness, whereas the replacement component gained importance. In environmentally homogeneous metacommunities, dispersal limitation created community dissimilarity via species replacement whereas the nestedness component remained low and unchanged across dispersal levels. Our study provides novel insights into how environmental heterogeneity and dispersal interact and shape metacommunity structure.

One contribution of 18 to a theme issue 'Human influences on evolution, and the ecological and societal consequences'. Urbanization causes both changes in community composition and evolutionary responses, but most studies focus on these responses in isolation. We performed an integrated analysis assessing the relative contribution of intra-and interspecific trait turnover to the observed change in zooplankton community body size in 83 cladoceran communities along urbanization gradients quantified at seven spatial scales (50-3200 m radii). We also performed a quantitative genetic analysis on 12 Daphnia magna populations along the same urbanization gradient. Body size in zooplankton communities generally declined with increasing urbanization, but the opposite was observed for communities dominated by large species. The contribution of intraspecific trait variation to community body size turnover with urbanization strongly varied with the spatial scale considered, and was highest for communities dominated by large cladoceran species and at intermediate spatial scales. Genotypic size at maturity was smaller for urban than for rural D. magna populations and for animals cultured at 248C compared with 208C. While local genetic adaptation likely contributed to the persistence of D. magna in the urban heat islands, buffering for the phenotypic shift to larger body sizes with increasing urbanization, community body size turnover was mainly driven by non-genetic intraspecific trait change.This article is part of the themed issue 'Human influences on evolution, and the ecological and societal consequences'.

As human population size increases and cities become denser, several urban-related selection pressures increasingly affect species composition in both terrestrial and aquatic habitats. Yet, it is not well known whether and how urbanization influences other facets of biodiversity, such as the functional and evolutionary composition of communities, and at what spatial scale urbanization acts. Here we used a hierarchical sampling design in which urbanization levels were quantified at seven spatial scales (ranging from 50 to 3200 m radii). We found that urbanization gradients are associated with a strong shift in cladoceran zooplankton species traits, which in turn affected phylogenetic composition of the entire metacommunity, but only when considering urbanization at the smallest spatial scale (50 m radius). Specifically, small cladoceran species dominated in more urbanized ponds whereas large-bodied, strong competitors prevailed in less urbanized systems. We also show that trait and phylogenetic metrics strongly increase the amount of variation in b-diversity that can be explained by degree of urbanization, environmental and spatial factors. This suggests that the mechanisms shaping b-diversity in our study system are mediated by traits and phylogenetic relatedness rather than species identities. Our study indicates that accounting for traits and phylogeny in metacommunity analyses helps to explain seemingly idiosyncratic patterns of variation in zooplankton species composition along urbanization gradients. The fact that urbanization acts only at the smallest spatial scale suggests that correctly managing environmental conditions locally has the power to counteract the effects of urbanization on biodiversity patterns. The multidimensional approach we applied here can be applied to other systems and organism groups and seems to be key in understanding how overall biodiversity changes in response to anthropogenic pressures and how this scales up to affect ecosystem functioning.

A negative consequence of biodiversity loss is reduced rates of ecosystem functions. Phylogenetic-based biodiversity indices have been claimed to provide more accurate predictions of ecosystem functioning than species diversity alone. This approach assumes that the most relevant traits for ecosystem functioning present a phylogenetic signal. Yet, traits-mediating niche partitioning and resource uptake efficiency in animals can be labile. To assess the relative power of a key trait (body size) and phylogeny to predict zooplankton top-down control on phytoplankton, we manipulated trait and phylogenetic distances independently in microcosms while holding species richness constant. We found that body size provided strong predictions of top-down control. In contrast, phylogeny was a poor predictor of grazing rates. Sizerelated grazing efficiency asymmetry was mechanistically more important than niche differences in mediating ecosystem function in our experimental settings. Our study demonstrates a strong link between a single functional trait (i.e. body size) in zooplankton and trophic interactions, and urges for a cautionary use of phylogenetic information and taxonomic diversity as substitutes for trait information to predict and understand ecosystem functions.

Biodiversity is structured by multiple mechanisms that are dependent, at least in part, on ecological similarities and differences among species. Integrating traits and phylogenies in diversity metrics may provide deeper insight into community assembly processes across spatial scales. However, different traits are influenced by processes at different spatial scales, and it is not clear how trait-spatial scale mismatches skew our ability to detect assembly patterns. An additional complexity is how phylogenetic distances, which might capture unmeasured traits, reflect spatially dependent processes. Here we analyze a freshwater zooplankton dataset from 91 ponds and show that different traits are associated with processes at different spatial scales. We first assessed the response of individual traits to processes at both α-and β-scales, and then quantified the power of different combinations of traits and phylogenetic distances to reveal environmental and spatial drivers of α-and β-diversity. We found that explanatory power was maximised when we accounted for environmental and spatial drivers with single, but different traits for α-and β-diversity. Using the most appropriate trait for each spatial scale outperformed phylogenetic information, but phylogenetic information outperformed the same traits when these were used at the wrong spatial scale, and all outperformed taxonomic analyses that ignore trait and phylogenetic information. We demonstrate that accounting for species' similarities and differences provides important information about dominant assembly mechanisms at different spatial scales, and that phylogeny is especially useful when measured traits are uninformative at a given spatial scale or when there is lack of trait data. Our study also indicates, however, that trait-scale mismatches among phylogenetically conserved traits may affect the performance of phylogenetic indices compared to indices that account only for the best single trait at each spatial scale.

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