Ecological communities are organized in complex ecological networks. Trait‐based analyses of the structure of these networks in highly diversified species assemblages are crucial for improving our understanding of the ecological and evolutionary processes causing specialization in mutualistic networks. In this study, we assessed the importance of morphological traits for structuring plant–hummingbird networks in Neotropical forests by using a novel combination of quantitative analytical approaches. We recorded the visitation of hummingbirds to plant species over an entire year at three different elevations in Costa Rica and constructed quantitative networks based on interaction frequencies. Three morphological traits were measured in hummingbirds (bill length, bill curvature, and body mass) and plants (corolla length, curvature, and volume). We tested the effects of avian morphological traits and abundance on ecological specialization of hummingbird species. All three morphological traits of hummingbirds were positively associated with ecological specialization, especially bill curvature. We tested whether interaction strength in the networks was associated with the degree of trait matching between corresponding pairs of morphological traits in plant and hummingbird species and explore whether this was related to resource handling times by hummingbird species. We found strong and significant associations between interaction strength and the degree of trait matching. Moreover, the degree of trait matching, particularly between bill and corolla length, was associated with the handling time of nectar resources by hummingbirds. Our findings show that bill morphology structures tropical plant–hummingbird networks and patterns of interactions are closely associated with morphological matches between plant and bird species and the efficiency of hummingbirds' resource use. These results are consistent with the findings of seminal studies in plant–hummingbird systems from the neotropics. We conclude that trait‐based analyses of quantitative networks contribute to a better mechanistic understanding of the causes of specialization in ecological networks and could be valuable for studying processes of complementary trait evolution in highly diversified species assemblages.

Knowledge of species composition and their interactions, in the form of interaction networks, is required to understand processes shaping their distribution over time and space. As such, comparing ecological networks along environmental gradients represents a promising new research avenue to understand the organization of life. Variation in the position and intensity of links within networks along environmental gradients may be driven by turnover in species composition, by variation in species abundances and by abiotic influences on species interactions. While investigating changes in species composition has a long tradition, so far only a limited number of studies have examined changes in species interactions between networks, often with differing approaches. Here, we review studies investigating variation in network structures along environmental gradients, highlighting how methodological decisions about standardization can influence their conclusions. Due to their complexity, variation among ecological networks is frequently studied using properties that summarize the distribution or topology of interactions such as number of links, connectance, or modularity. These properties can either be compared directly or using a procedure of standardization. While measures of network structure can be directly related to changes along environmental gradients, standardization is frequently used to facilitate interpretation of variation in network properties by controlling for some co-variables, or via null models. Null models allow comparing the deviation of empirical networks from random expectations and are expected to provide a more mechanistic understanding of the factors shaping ecological networks when they are coupled with functional traits. As an illustration, we compare approaches to quantify the role of trait matching in driving the structure of plant-hummingbird mutualistic networks, i.e. a direct comparison, standardized by null models and hypothesis-based metaweb. Overall, our analysis warns against a comparison of studies that rely on distinct forms of standardization, as they are likely to highlight different signals. Fostering a better understanding of the analytical tools available and the signal they detect will help produce deeper insights into how and why ecological networks vary along environmental gradients.