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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.

The conversion of tropical habitats has dramatic implications on biodiversity and represents one of the greatest conservation challenges of our time. Seasonally Dry Tropical Forests (SDTF), which are disjointly distributed throughout the Neotropics, are especially susceptible to human activities. The Caatinga Dry Forest, located in the semi-arid interior of northeastern Brazil, represents not only the largest and most biologically diverse nucleus of SDTF, but also the world’s most densely populated semi-arid region, with ever-growing pressure on its natural resources. To prevent illegal logging, conservation agencies looked at forest management, where an area is divided in smaller stands which are gradually logged and allowed to regrow for a period of time, when a new cutting cycle should reinitiate. The impacts of these management schemes on biodiversity, however, remain largely untested. We conducted standardized avian surveys to evaluate the effects of forest management on the avian community at a 1,670 ha privately owned property located on the Chapada do Araripe, northeastern Brazil. This area was divided in 22 forest stands, half of which had already been logged at the time of our sampling, creating a gradient of logged and natural forests and an 11-yr chrono-sequence of forest regeneration. Our results show that logged areas present fewer individuals, fewer species, and different avian assemblages than unlogged forests. Such differences are mostly driven by forest-dependent species, which were overwhelmingly affected by forest management. Our results show that although logged forests tend to recover its height after a decade, they do not recover the originally forest cover, measured by the Normalized Difference Vegetation Index. Likewise, decade-long recovering stands continue to show lower species richness, lower bird abundance, and different avian composition than unlogged forests. We identified a set of bird species that are more affected by forest management (ecological losers) and a group of birds that apparently benefit from the referred changes in land use (ecological winners). We conclude that completely managing an entire area may cause the extirpation of several forest-dependent species. We therefore suggest keeping logged and unlogged plots intermingled, to avoid local extinctions and the complete modification of the original avifauna.

Author(s): Araujo, AC; Martin Gonzalez, AM; Sandel, B; Maruyama, PK; Fischer, E; Vizentin-Bugoni, J; de Araujo, FP; Coelho, AG; Faria, RR; Kohler, G; Las-Casas, FMG; Lopes, AV; Machado, AO; Machado, CG; Machado, IC; McGuire, JA; Moura, AC; Oliveira, GM; Oliveira, PE; Rocca, MA; Rodrigues, LDC; Rodrigues, M; Rui, AM; Sazima, I; Sazima, M; Varassin, IG; Wang, Z; Dalsgaard, B; Svenning, JC | Abstract: 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.

Ecological communities that experience stable climate conditions have been speculated to preserve more specialized interspecific associations and have higher proportions of smaller ranged species (SRS). Thus, areas with disproportionally large numbers of SRS are expected to coincide geographically with a high degree of community-level ecological specialization, but this suggestion remains poorly supported with empirical evidence. Here, we analysed data for hummingbird resource specialization, range size, contemporary climate, and Late Quaternary climate stability for 46 hummingbird–plant mutualistic networks distributed across the Americas, representing 130 hummingbird species ( ca 40% of all hummingbird species). We demonstrate a positive relationship between the proportion of SRS of hummingbirds and community-level specialization, i.e. the division of the floral niche among coexisting hummingbird species. This relationship remained strong even when accounting for climate, furthermore, the effect of SRS on specialization was far stronger than the effect of specialization on SRS, suggesting that climate largely influences specialization through species' range-size dynamics. Irrespective of the exact mechanism involved, our results indicate that communities consisting of higher proportions of SRS may be vulnerable to disturbance not only because of their small geographical ranges, but also because of their high degree of specialization.