Interactions between species are influenced by different ecological mechanisms, such as morphological matching, phenological overlap and species abundances. How these mechanisms explain interaction frequencies across environmental gradients remains poorly understood. Consequently, we also know little about the mechanisms that drive the geographical patterns in network structure, such as complementary specialization and modularity. Here, we use data on morphologies, phenologies and abundances to explain interaction frequencies between hummingbirds and plants at a large geographical scale. For 24 quantitative networks sampled throughout the Americas, we found that the tendency of species to interact with morphologically matching partners contributed to specialized and modular network structures. Morphological matching best explained interaction frequencies in networks found closer to the equator and in areas with low-temperature seasonality. When comparing the three ecological mechanisms within networks, we found that both morphological matching and phenological overlap generally outperformed abundances in the explanation of interaction frequencies. Together, these findings provide insights into the ecological mechanisms that underlie geographical patterns in resource specialization. Notably, our results highlight morphological constraints on interactions as a potential explanation for increasing resource specialization towards lower latitudes.

Aim Species interaction networks are known to vary in structure over large spatial scales. We investigated the hypothesis that environmental factors affect interaction network structure by influencing the functional diversity of ecological communities. Notably, we expect more functionally diverse communities to form interaction networks with a higher degree of niche partitioning. Location: Americas. Time period: Current. Major taxa studied: Hummingbirds and their nectar plants. Methods We used a large dataset comprising 74 quantitative plant–hummingbird interaction networks distributed across the Americas, along with morphological trait data for 158 hummingbird species. First, we used a model selection approach to evaluate associations between the environment (climate, topography and insularity), species richness and hummingbird functional diversity as predictors of network structure (niche partitioning, i.e., complementary specialization and modularity). Second, we used structural equation models (SEMs) to ask whether environmental predictors and species richness affect network structure directly and/or indirectly through their influence on hummingbird functional diversity. For a subset of 28 networks, we additionally evaluated whether plant functional diversity was associated with hummingbird functional diversity and network structure. Results Precipitation, insularity and plant richness, together with hummingbird functional diversity (specifically, functional dispersion), were consistently strong predictors of niche partitioning in plant–hummingbird networks. Moreover, SEMs showed that environmental predictors and plant richness affected network structure both directly and indirectly through their effects on hummingbird functional diversity. Plant functional diversity, however, was unrelated to hummingbird functional diversity and network structure. Main conclusions: We reveal the importance of hummingbird functional diversity for niche partitioning in plant–hummingbird interaction networks. The lack of support for similar effects for plant functional diversity potentially indicates that consumer functional diversity might be more important for structuring interaction networks than resource functional diversity. Changes in pollinator functional diversity are therefore likely to alter the structure of interaction networks and associated ecosystem functions.

Functional traits can determine pairwise species interactions, such as those between plants and pollinators. However, the effects of biogeography and evolutionary history on trait‐matching and trait‐mediated resource specialization remain poorly understood. We compiled a database of 93 mutualistic hummingbird–plant networks (including 181 hummingbird and 1,256 plant species), complemented by morphological measures of hummingbird bill and floral corolla length. We divided the hummingbirds into their principal clades and used knowledge on hummingbird biogeography to divide the networks into four biogeographical regions: Lowland South America, Andes, North & Central America, and the Caribbean islands. We then tested: (a) whether hummingbird clades and biogeographical regions differ in hummingbird bill length, corolla length of visited flowers and resource specialization, and (b) whether hummingbirds' bill length correlates with the corolla length of their food plants and with their level of resource specialization. Hummingbird clades dominated by long‐billed species generally visited longer flowers and were the most exclusive in their resource use. Bill and corolla length and the degree of resource specialization were similar across mainland regions, but the Caribbean islands had shorter flowers and hummingbirds with more generalized interaction niches. Bill and corolla length correlated in all regions and most clades, that is, trait‐matching was a recurrent phenomenon in hummingbird–plant associations. In contrast, bill length did not generally mediate resource specialization, as bill length was only weakly correlated with resource specialization within one hummingbird clade (Brilliants) and in the regions of Lowland South America and the Andes in which plants and hummingbirds have a long co‐evolutionary history. Supplementary analyses including bill curvature confirmed that bill morphology (length and curvature) does not in general predict resource specialization. These results demonstrate how biogeographical and evolutionary histories can modulate the effects of functional traits on species interactions, and that traits better predict functional groups of interaction partners (i.e. trait‐matching) than resource specialization. These findings reveal that functional traits have great potential, but also key limitations, as a tool for developing more mechanistic approaches in community ecology. A free Plain Language Summary can be found within the Supporting Information of this article.

Abundant pollinators are often more generalised than rare pollinators. This could be because abundant species have more chance encounters with potential interaction partners. On the other hand, generalised species could have a competitive advantage over specialists, leading to higher abundance. Determining the direction of the abundance–generalisation relationship is therefore a ‘chicken‐and‐egg’ dilemma. Here we determine the direction of the relationship between abundance and generalisation in plant–hummingbird pollination networks across the Americas. We find evidence that hummingbird pollinators are generalised because they are abundant, and little evidence that hummingbirds are abundant because they are generalised. Additionally, most patterns of species‐level abundance and generalisation were well explained by a null model that assumed interaction neutrality (interaction probabilities defined by species relative abundances). These results suggest that neutral processes play a key role in driving broad patterns of generalisation in animal pollinators across large spatial scales.

Aim We examined the effects of space, climate, phylogeny and species traits on module composition in a cross‐biomes plant–hummingbird network. Location Brazil, except Amazonian region. Methods We compiled 31 local binary plant–hummingbird networks, combining them into one cross‐biomes metanetwork. We conducted a modularity analysis and tested the relationship between species’ module membership with traits, geographical location, climatic conditions and range sizes, employing random forest models. We fitted reduced models containing groups of related variables (climatic, spatial, phylogenetic, traits) and combinations of groups to partition the variance explained by these sets into unique and shared components. Results The Brazilian cross‐biomes network was composed of 479 plant and 42 hummingbird species, and showed significant modularity. The resulting six modules conformed well to vegetation domains. Only plant traits, not hummingbird traits, differed between modules, notably plants’ growth form, corolla length, flower shape and colour. Some modules included plant species with very restricted distributions, whereas others encompassed more widespread ones. Widespread hummingbirds were the most connected, both within and between modules, whereas widespread plants were the most connected between modules. Among traits, only nectar concentration had a weak effect on among‐module connectivity. Main conclusions Climate and spatial filters were the main determinants of module composition for hummingbirds and plants, potentially related to resource seasonality, especially for hummingbirds. Historical dispersal‐linked contingency, or environmental variations not accounted for by the explanatory factors here evaluated, could also contribute to the spatial component. Phylogeny and morphological traits had no unique effects on the assignment of species to modules. Widespread species showed higher within‐ and/or among‐module connectivity, indicating their key role connecting biomes, and, in the case of hummingbirds, communities within biomes. Our results indicate that biogeography and climate not only determine the variation of modularity in local plant–animal networks, as previously shown, but also affect the cross‐biomes network structure.