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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.

Indirect interactions are central to ecological and evolutionary dynamics in pollination communities, yet we have little understanding about the processes determining patterns of indirect interactions, such as those between pollinators through shared flowering plants. Instead, research has concentrated on the processes responsible for direct interactions and whole-network structures. This is partly due to a lack of appropriate tools for characterising indirect interaction structures, because traditional network metrics discard much of this information. The recent development of tools for counting motifs (subnetworks depicting interactions between a small number of species) in bipartite networks enables detailed analysis of indirect interaction patterns. Here we generate plant–hummingbird pollination networks based on three major assembly processes—neutral effects (species interacting in proportion to abundance), morphological matching and phenological overlap—and evaluate the motifs associated with each one. We find that different processes produce networks with significantly different patterns of indirect interactions. Neutral effects tend to produce densely connected motifs, with short indirect interaction chains, and motifs where many specialists interact indirectly through a single generalist. Conversely, niche-based processes (morphology and phenology) produced motifs with a core of interacting generalists, supported by peripheral specialists. These results have important implications for understanding the processes determining indirect interaction structures. A free Plain Language Summary can be found within the Supporting Information of this article.

Anthropogenic disturbances decrease the resources available to ants and simplify the ant fauna. This could affect ant–plant interactions, especially if key-species disappear. In terms of conservation, disturbances can select for ants resilient to disturbances, and such species can be considered indicators of succession processes. Here, we investigated the extent to which a vegetation clearcutting event (by bulldozer) affected the communities of both plants (seven species of Bignoniaceae) and associated ants. We hypothesized that only generalist ants would remain after clearcutting. Clearcutting reduced plant abundance, with only 40% of individuals resprouting after disturbance. Plant species richness was not affected by clearcutting, but ant species richness was lower in disturbed than in undisturbed conditions (four instead of seven species). Camponotus ants were dominant and were found regularly on all plant species before and after clearcutting. These ants had high centrality values and were regarded as influential species in the structure of the ant–plant network. Camponotus are generalists and nest in the soil; thus, clearcutting did not affect them to the same degree as the ants that nest in plants. The ant–plant network showed no changes in the specialization index (H2′), indicating that even with losses of some ant species, the network properties were maintained: plants were still being visited by ants. Therefore, we show that clearcutting had a weak effect on ant–plant interactions, especially because Camponotus buffered the losses of other ants.

Animal pollinators are globally threatened by anthropogenic land use change and agricultural intensification. The yield of many food crops is therefore negatively impacted because they benefit from biotic pollination. This is especially the case in the tropics. For instance, fruit set of Coffea arabica has been shown to increase by 10–30% in plantations with a high richness of bee species, possibly influenced by the availability of surrounding forest habitat. Here, we performed a global literature review to (1) assess how much animal pollination enhances coffee fruit set, and to (2) examine the importance of the amount of forest cover, distance to nearby forest and forest canopy density for bee species richness and coffee fruit set. Using a systematic literature review, we identified eleven case studies with a total of 182 samples where fruit set of C. arabica was assessed. We subsequently gathered forest data for all study sites from satellite imagery. We modelled the effects of open (all forest with a canopy density of ≥25%), closed (≥50%) and dense (≥75%) forests on pollinator richness and fruit set of coffee. Overall, we found that animal pollination increases coffee fruit set by ~18% on average. In only one of the case studies, regression results indicate a positive effect of dense forest on coffee fruit set, which increased with higher forest cover and shorter distance to the forest. Against expectations, forest cover and distance to open forest were not related to bee species richness and fruit set. In summary, we provide strong empirical support for the notion that animal pollinators increase coffee fruit set. Forest proximity had little overall influence on bee richness and coffee fruit set, except when farms were surrounded by dense tropical forests, potentially because these may provide high-quality habitats for bees pollinating coffee. We, therefore, advocate that more research is done to understand the biodiversity value of dense forest for pollinators, notably assessing the mechanisms underlying the importance of forest for pollinators and their pollination services.

Cities harbour considerable biodiversity and there has been an increased concern about the conservation of pollinators in urban environments. Here, we evaluated how urbanization affects plant-hummingbird interactions at two spatial scales. First, in a medium-sized city from southeastern Brazil (>600,000 inhabitants), we contrasted interaction networks from urban and natural areas, and used artificial nectar feeder stations to evaluate changes in the composition of hummingbird assemblages across an urbanization gradient. Second, we compiled data on six urban plant-hummingbird interaction networks from south and southeastern Brazil to identify the characteristics associated with the most important plants. Locally, urbanization affected hummingbird communities by promoting higher generalization and dominance by more aggressive hummingbirds. Notably, specialized long-billed hermits were absent both in the urban interaction network and at feeder stations from more urbanized areas. Across networks, trees were more important for hummingbirds than shrubs/herbs as were specialized ornithophilous flowers in relation to non-ornithophilous flowers. Plant origin (native or exotic) did not matter. Our results indicate that urban plant-hummingbird communities are organized differently than their counterparts from natural areas, which usually feature key hermits and few trees. Since hermits provide important pollination services, especially for specialized ornithophilous plants, initiatives such as green corridors and preference for native plants with specialized hummingbird-pollinated flowers in urban landscaping may contribute to community restoration and ecosystem functioning.

When describing plant-animal interaction networks, sampling can be performed using plant- or animal-centred approaches. Despite known effects of sampling on network structure, how samplings affect the estimates of interaction β-diversity across networks is still unresolved. We investigated how the sampling method affects the assessment of β-diversity of interactions, turnover and rewiring. We contrasted plant- and animal-centred sampling methods applied to pollination networks across habitats in a heterogeneous tropical landscape, the Pantanal Wetlands. We also asked whether plant traits influence the difference in interaction specialization according to sampling. Plant-centred networks resulted in higher β-diversity of interactions in space than animal-centred networks. Turnover explained most of the β-diversity in both methods, but rewiring was proportionately more important when using the animal-centred method. While the plant-centred method indicated lower network modularity and specialization, floral traits modulated the effects of the sampling method on species-level network metrics. Combining animal- and plant-centred approaches returned intermediate values for β-diversity of interactions and network metrics. Distinct methods may also be better suited for answering questions at different scales. Our results point out that the method choice, or combination of methods, should always reflect the appropriate scale of the factors determining the interactions being investigated.

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.

Made available in DSpace on 2018-11-26T17:55:26Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-07-01. Added 1 bitstream(s) on 2019-10-09T18:27:51Z : No. of bitstreams: 1 S0102-33062018000300386.pdf: 1047685 bytes, checksum: 57b1bc1e74dcec638d108f4a395975d7 (MD5) Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Most angiosperms rely on animals for pollination, and insects, especially bees, are the most frequent pollinators. Many native Neotropical plants are frequently visited by the invasive honeybee (Apis mellifera), but its role in the pollination of these plants has been little investigated. We assessed the contribution of various floral visitors, including native bees and the honeybee, on the pollination of a generalist rupestrian grassland daisy, Aspilia jolyana (Asteraceae), in Serra do Cipo, Espinhaco Mountain Range, Brazil. We recorded floral visitors and measured the seed set resulting from one single visitation. We observed a total of 442 visits, mostly by bees, with Bombus pauloensis and Apis mellifera being the most common floral visitors. Other visitors included many other species of bees, flies, hummingbirds, wasps and butterflies. Pollinators significantly increased seed set in comparison to non-visited (bagged) capitula. Moreover, there was no difference among bee species/groups in their contribution to seed set. Thus, A. jolyana benefits from its generalized pollination strategy, and frequent bee visitors, including several native species and the invasive honeybee, are equally effective pollinators for this generalist daisy of rupestrian grasslands. Univ Estadual Campinas, Inst Biol, Dept Biol Vegetal, BR-13083970 Campinas, SP, Brazil Univ Fed Uberlandia, Inst Biol, BR-38405302 Uberlandia, MG, Brazil Univ Illinois Champaign Urbana, Nat Resources & Environm Sci Dept, Urbana, IL 61820 USA Univ Copenhagen, Ctr Macroecol Evolut & Climate, Nat Hist Museum Denmark, DK-2100 Copenhagen, Denmark Univ Estadual Paulista, Inst Biociencias, Dept Bot, Lab Fenol, BR-13506900 Rio Claro, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Dept Bot, Lab Fenol, BR-13506900 Rio Claro, SP, Brazil CNPq: 400717/2013-1 FAPESP: 2013/50155-0 FAPESP: 2015/21457-4

The temporal dynamics of plant phenology and pollinator abundance across seasons should influence the structure of plant–pollinator interaction networks. Nevertheless, such dynamics are seldom considered, especially for diverse tropical networks. Here, we evaluated the temporal variation of four plant–pollinator networks in two seasonal ecosystems in Central Brazil (Cerrado and Pantanal). Data were gathered on a monthly basis over 1 year for each network. We characterized seasonal and temporal shifts in plant–pollinator interactions, using temporally discrete networks. We predicted that the greater floral availability in the rainy season would allow for finer partitioning of the floral niche by the pollinators, i.e. higher specialization patterns as previously described across large spatial gradients. Finally, we also evaluated how sampling restricted to peak flowering period may affect the characterization of the networks. Contrary to our expectations, we found that dry season networks, although characterized by lower floral resource richness and abundance, showed higher levels of network‐wide interaction partitioning (complementary specialization and modularity). For nestedness, though, this between‐seasons difference was not consistent. Reduced resource availability in the dry season may promote higher interspecific competition among pollinators leading to reduced niche overlap, thus explaining the increase in specialization. There were no consistent differences between seasons in species‐level indices, indicating that higher network level specialization is an emergent property only seen when considering the entire network. However, bees presented higher values of specialization and species strength in relation to other groups such as flies and wasps, suggesting that some plant species frequently associated with bees are used only by this group. Our study also indicates that targeted data collection during peak flowering generates higher estimates of network specialization, possibly because species activity spans longer periods than the targeted time frame. Hence, depending on the period of data collection, different structural values for the networks of interactions may be found. Synthesis. Plant–pollinator networks from tropical environments have structural properties that vary according to seasons, which should be taken into account in the description of the complex systems of interactions between plants and their pollinators in these areas.

Habitat complexity is an important factor structuring macroinvertebrate communities on macrophytes, and epiphytic hydroids may offer additional habitats for the epifauna. Caprellids are known to benefit from the association with hydroids, but it is unclear if epiphytic hydroids affect their abundance. In this study, we investigated if caprellid abundance is related to epiphytic hydroid abundance on Sargassum cymosum, and whether this is related to caprellids’ clinging behavior. We found that hydroid abundance had a positive effect on caprellid abundance, irrespective of species, sex, developmental and reproductive stages of caprellids, with the exception of Caprella scaura that present maternal care behavior. Experiments also showed that caprellids are more abundant on fronds of Sargassum with hydroids. Moreover, juvenile caprellids prefer to cling directly to hydroids and also prefer portions of the alga with hydroids attached. Our results indicate that epiphytic hydroids add structural complexity to macroalgae, favoring the occurrence of caprellids. However, this effect may be conditioned by the reproductive behavior of caprellid species, since hydroids are particularly important for juveniles by offering suitable structures for their small pereopods to grasp. This, in turn, may indirectly benefit adult caprellids, contributing to their choice of habitats that favor the performance of their juveniles.

Plant species within communities may overlap in pollinators' use and influence visitation patterns of shared pollinators, potentially engaging in indirect interactions (e.g., facilitation or competition). While several studies have explored the mechanisms regulating insect-pollination networks, there is a lack of studies on bird-pollination systems, particularly in species-rich tropical areas. Here, we evaluated if phenotypic similarity, resource availability (floral abundance), evolutionary relatedness and flowering phenology affect the potential for indirect effects via shared pollinators in hummingbird-pollinated plant species within four communities in the Brazilian Atlantic forest. Among the evaluated factors, phenotypic similarity (corolla length and anther height) was the most important variable, while resource availability (floral abundance) had a secondary importance. On the other hand, evolutionary relatedness and flowering phenology were less important, which altogether highlights the relevance of convergent evolution and that the contribution of a plant to the diet of the pollinators of another plant is independent of the level of temporal overlap in flowering in this tropical system. Interestingly, our findings contrast with results from multiple insect-pollinated plant communities, mostly from temperate regions, in which floral abundance was the most important driver, followed by evolutionary relatedness and phenotypic similarity. We propose that these contrasting results are due to high level of specialization inherent to tropical hummingbird-pollination systems. Moreover, our results demonstrated that factors defining linkage rules of plant-hummingbird networks also determinate plant-plant potential indirect effects. Future studies are needed to test if these findings can be generalized to other highly specialized systems. Overall, our results have important implications for the understanding of ecological processes due resource sharing in mutualistic systems.

Ecological network approaches may contribute to conservation practices by quantifying within-community importance of species. In mutualistic plant-pollinator systems, such networks reflect potential pollination of the plants and a considerable portion of the energy consumption by the pollinators, two key components for each party. Here, we used two different sampling approaches to describe mutualistic plant-hummingbird networks from a cloud forest in the Colombian Western Andes, home to the Colorful Puffleg Eriocnemis mirabilis, an endemic and critically endangered hummingbird. We contrast networks between two localities (a protected area inside a National park vs. its buffer zone) and across sampling methods (floral visitation vs. pollen loads) to assess how the network structure and the importance of each hummingbird species within the networks may change. Visitation networks were characterized as having higher sampling completeness, yet pollen load network recorded more pollen types than plant species recorded by visitation. Irrespective of the sampling methods, the Colorful Puffleg was one of the most important hummingbird species in the network within the protected area inside the National park, but not in the buffer zone. Moreover, most species-level network indices were related to hummingbirds’ abundance. This suggests that conservation initiatives aimed at the endangered Colorful Puffleg may both help on the survival of this endangered hummingbird, as well as on maintaining its key role in the mutualistic interaction network inside the National Park. Our study illustrates how conservation practitioners could assess the local importance of endangered species using interaction network approaches.

International audience; Species are entangled within communities by their interactions in such a manner that their local extinction may unchain coextinction cascades and impact community dynamics and stability. Despite increasing attention, simulation models to estimate the robustness of interaction networks largely neglect the important role of interaction rewiring, that is, the ability of species to switch partners. Here we propose a new method to incorporate the potential of species to replace lost partners into a widely used coextinction model to estimate network robustness. In this model, species are allowed to rewire their interactions after initial loss of partners according to probabilities derived from well‐known mechanisms that determine mutualistic interactions, for example, trait matching, phenological overlap and abundances. To illustrate the use of this method, we analyzed a well‐sampled dataset of mutualistic plant–hummingbird interactions. We found that (a) in general, rewiring increases the estimated robustness, (b) networks are similarly robust to the loss of pollinators or plants, (c) morphological matching and phenological overlap leads to higher robustness, (d) when multiple rewiring mechanisms are combined, however, robustness increases little, and (e) rewiring tends to increase robustness more in scenarios when the loss of generalist species occurs first. Our results suggest that the same mechanisms known to drive plant–hummingbird network structure are relevant in buffering the effects of species loss via rewiring. The method proposed here can be applied to a wide range of interaction networks and is flexible to the inclusion of other variables important in determining interactions for specific systems. It also allows changes on the assumptions regarding the importance of distinct mechanisms, for instance including a hierarchical relationship, which facilitates insights into the relative importance of multiple variables influencing network disassembling. The analytical framework we offer here represents a step towards a more realistic estimation on how species loss may affect the integrity of interaction networks.

Abundant pollinators are often more generalised than rare pollinators. This could be because abundance drives generalisation: neutral effects suggest that more abundant species will be more generalised simply because they have more chance encounters with potential interaction partners. On the other hand, generalisation could drive abundance, as generalised species could have a competitive advantage over specialists, being able to exploit a wider range of resources and gain a more balanced nutrient intake. 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 sampled from a variety of locations across the Americas. For the first time we resolve the direction of the abundance-generalisation relationship using independent data on animal abundance. We find evidence that hummingbird pollinators are generalised because they are abundant, and little evidence that hummingbirds are abundant because they are generalised. Additionally, a null model analysis suggests this pattern is due to neutral processes: most patterns of species-level abundance and generalisation were well explained by a null model that assumed interaction neutrality. These results suggest that neutral processes play a key role in driving broad patterns of generalisation in animal pollinators across large spatial scales.DeclarationsFunding – BIS is supported by the Natural Environment Research Council as part of the Cambridge Earth System Science NERC DTP [NE/L002507/1]. JVB was funded by CERL - Engineer Research and Development Center. PKM was funded by the São Paulo Research Foundation (FAPESP grant #2015/21457-4). PAC was funded by the David Lack studentship from the British Ornithologists’ Union and Wolfson College, University of Oxford. CL was funded by the ESDEPED-UAT grant. MAM acknowledges the Consejo Nacional para Investigaciones Científicas y Tecnológicas (Costa Rica), German Academic Exchange Service and the research funding program ‘LOEWE-Landes-Offensive zur Entwicklung Wissenschaftlichö konomischer Exzellenz’ of Hesse’s Ministry of Higher Education, Research, and the Arts (Germany). ROP was funded by CONACyT (project 258364). MAR was supported by the State of São Paulo Research Foundation (FAPESP) within the BIOTA/FAPESP, The Biodiversity Institute Program (www.biota.org.br) and the ‘Parcelas Permanentes’ project, as well as by Coordenação de Pessoal de Nível Superior (CAPES), Fundo de Apoio ao Ensino e à Pesquisa (FAEP)/Funcamp/Unicamp and The Nature Conservancy (TNC) of Brazil. LCR was supported by CNPq and Capes. MS was funded by CNPq (grant #302781/2016-1). AMMG is supported through a Marie Skłodowska-Curie Individual Fellowship (H2020-MSCA-IF-2016-704409). LVD was supported by the Natural Environment Research Council (grants NE/K015419/1 and NE/N014472/1). AMMG, JS, CR and BD thank the Danish National Research Foundation for its support of the Center for Macroecology, Evolution and Climate (grant no. DNRF96). WJS is funded by Arcadia.

• Floral visitors differ in their efficacy as pollinators and the impact of different pollinator species on the pollen flow and plant reproduction has been frequently evaluated. In contrast, the impact of within-species foraging changes on their efficacy as pollinators has been seldom quantified. • We studied a self-incompatible shrub Palicourea rigida (Rubiaceae) and its hummingbird pollinators which adjust their behaviour according to floral resources availability. Fluorescence microscopy was used to access pollen tube growth and incompatibility reaction in pistils after a single visit of territorial or intruder hummingbirds in two populations. To characterize the plant populations and possible differences on resource availability between areas we used a three-term quadrat variance method to detect clusters of floral resources. • Within species variation in foraging behaviour, but not the species identity affected pollinator efficacy. Effectively, hummingbirds intruding territories deposited more compatible pollen grains on P. rigida stigmas than territory holders in both study areas. Additionally, territory holders deposited more incompatible than compatible pollen grains. • Our results imply that intraspecific foraging behaviour variation has consequences for pollination success. Quantifying such variation and addressing the implications of intraspecific variability contribute to a better understanding of the dynamics and consequences of plant-pollinators interactions. This article is protected by copyright. All rights reserved.

Hummingbirds are the most common group of vertebrate pollinators in the Neotropics, associated with pollination of ornithophilous flowers. The group can be divided into hermits and non-hermits, in which hermits are often regarded as the most specialized group closely associated with more specialized ornithophilous plants. In this study, we investigated the association of ornithophilous flowers and hummingbirds in two small fragments of forest surrounded by an urban matrix. We characterized the species composition, abundance and phenology of the hummingbirds and ornithophilous plants and related pollinator abundance to floral resource availability. Our results were compared to published data from other forest areas from the Cerrado. In our study areas, the diversity of ornithophilous plants and hummingbirds was similar to what has been reported for forest fragments in the Cerrado. Nevertheless, we found Thalurania furcata, a non-hermit, acting as the most frequent hummingbird pollinator in contrast to more preserved areas in which a hermit, Phaethornis pretrei, is commonly found as the main pollinator for ornithophilous plants. The nectar energy availability at the plant population level was the only factor associated with hummingbird visitation rates, suggesting that a higher availability of nectar resources in the fragments attracts greater abundance of birds. The unusual setting of having a non-hermit species as the main hummingbird pollinator in forest areas suggests that habitat fragmentation can favor more generalist hummingbird species, and this potentially has consequence for the pollination of associated plants.

PLANT-POLLINATOR INTERACTIONS IN HIGHLAND VEGETATION IN THE ATLANTIC FOREST Tropical high-altitude vegetation is unique due to susceptibility to severe weather conditions in relation to lower for­mations, and by the peculiarity of its flora with many relictual components. Studies on plant-pollinator interactions in high-altitude rocky outcrops and forests of the Atlantic Forest are scarce, but compilation of information allows us to identify some patterns: low frequency of visits, high floral longevity and generalized pollination system. In tropical mountain ecosystems, the degree of generalization of pollination systems in functional (pollinator groups) and ecological (number of species) terms tends to be high, mainly due to the over-representation of certain plant taxa ( e.g . , Asteraceae in rocky outcrops and Fabaceae, Myrtaceae, Rubiaceae and Sapindaceae in montane forests). Generalized pollination systems and autogamy may be advantageous for tropical high-altitude plants due to the more severe weather conditions ( e.g. , low temperature), which decrease abundance and limit the activity of pollinators, resulting in lower visitation frequency. Nevertheless, some well represented groups in forests, such as orchids and plants pollinated by hummingbirds and bats, exemplify cases of higher functional specialization, as well as plants with poricidal anthers pollinated by bees in the high-altitude grasslands. However, in rocky outcrops, for some functional groups of pollinators ( e.g ., hummingbirds, bats, beetles and hawkmoths), the availability of resources does not allow the maintenance of all species throughout the year, favoring possible local or altitudinal migrations. Thus, rocky outcrops and high-altitude forests constitute a unit in the sense of sustaining the pollinator community. Indeed ro­cky outcrops and high-altitude forests share an evolutionary history at the regional scale since they passed through similar events of expansion and retraction in response to climate changes in the Quaternary. This could explain the complementarity between the two types of vegetation in the use of floral resources by pollinators. Besides the associations identified here, the ecology and evolution of plant-pollinator interactions in high-altitude vegetation of the Atlantic Forest remain poorly understood, making urgent the development of an integrative research program, as well as projects on issues related to climate change and biodiversity conservation.

Elucidating the factors determining the occurrence of florivorous organisms is an essential step for comprehending arthropod–plant interactions, especially when considering florivores that use flowers/inflorescences as microhabitats. In this study, we characterize the interaction between florivorous thrips (Thysanoptera) and Palicourea rigida (Rubiaceae), a distylous hummingbird-pollinated shrub. We investigated the relative role of different factors in determining thrips occurrence in the flower and inflorescence microhabitats. Furthermore, we experimentally examined the protective role of corolla influencing thrips exploration of floral buds. Frankliniella musaeperda (Thripidae) was the only species recorded on P. rigida, feeding on floral tissue, pollen and nectar. Thrips occurrence was not related to distyly, but rather to floral stage. Open flowers presented the highest abundance of thrips, followed by senescent flowers and then buds. The experimental opening of buds translated in increased thrips occurrence, indicating that F. musaeperda manage to explore the microhabitat offered by the floral chamber, as long as there is an opening in the corolla. In inflorescences, thrips abundance was negatively related to the number of ants visiting extrafloral nectaries. We found that the marked difference between floral morphs of distylous plants is not necessarily reflected in the abundance of florivores. Thrips seek for floral cavities, preferentially those with fresh tissue, which may confer nutrient-rich food and protection. Buds also provide this; however, the enclosed petals are an effective barrier against F. musaeperda entrance. At inflorescence scale, presence of mutualistic ants in high numbers can drive away these flower-feeding insects. Despite the abundance of thrips in the flowers, there was no evidence of any functional relationship, either of pollination for flowers or of breeding for insects. We demonstrate here that in the flower/inflorescence microhabitat, structural and biotic factors play a key role in the exploitation and occupation by insect florivores.

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.

Most tropical plants rely on animals for pollination, thus engaging in complex interaction networks. Here, we present a global overview of pollination networks and point out research gaps and emerging differences between tropical and non-tropical areas. Our review highlights an uneven global distribution of studies biased towards non-tropical areas. Moreover, within the tropics, there is a bias towards the Neotropical region where partial networks represent 70.1% of the published studies. Additionally, most networks sampled so far (95.6%) were assembled by inferring interactions by surveying plants (a phytocentric approach). These biases may limit accurate global comparisons of the structure and dynamics of tropical and non-tropical pollination networks. Noteworthy differences of tropical networks (in comparison to the non-tropical ones) include higher species richness which, in turn, promotes lower connectance but higher modularity due to both the higher diversity as well as the integration of more vertebrate pollinators. These interaction patterns are influenced by several ecological, evolutionary, and historical processes, and also sampling artifacts. We propose a neutral–niche continuum model for interactions in pollination systems. This is, arguably, supported by evidence that a high diversity of functional traits promotes greater importance of niche-based processes (i.e., forbidden links caused by morphological mismatching and phenological non-overlap) in determining which interactions occur, rather than random chance of encounter based on abundances (neutrality). We conclude by discussing the possible existence and direction of a latitudinal gradient of specialization in pollination networks.

Convergent reproductive traits in non-related plants may be the result of similar environmental conditions and/or specialised interactions with pollinators. Here, we documented the pollination and reproductive biology of Bionia coriacea (Fabaceae), Esterhazya splendida (Orobanchaceae) and Ananas ananassoides (Bromeliaceae) as case studies in the context of hummingbird pollination in Cerrado, the Neotropical savanna of Central South America. We combined our results with a survey of hummingbird pollination studies in the region to investigate the recently suggested association of hummingbird pollination and self-compatibility. Plant species studied here differed in their specialisation for ornithophily, from more generalist A. ananassoides to somewhat specialist B. coriacea and E. splendida. This continuum of specialisation in floral traits also translated into floral visitor composition. Amazilia fimbriata was the most frequent pollinator for all species, and the differences in floral display and nectar energy availability among plant species affect hummingbirds' behaviour. Most of the hummingbird-pollinated Cerrado plants (60.0%, n = 20), including those studied here, were self-incompatible, in contrast to other biomes in the Neotropics. Association to more generalist, often territorial, hummingbirds, and resulting reduced pollen flow in open savanna areas may explain predominance of self-incompatibility. But it is possible that mating system is more associated with the predominance of woody hummingbird plants in the Cerrado plant assemblage than to the pollination system itself.

Providing hummingbirds with artificial feeders containing sugar solution is common practice throughout the Americas. Although feeders can affect hummingbird foraging behavior and abundance, it is poorly understood how far this effect may extend. Moreover, it remains debated whether nectar-feeders have a negative impact on hummingbird-pollinated plants by reducing flower visitation rates and pollen transfer close to the feeders. Here, we investigated the effects of distance to nectar-feeders on a local hummingbird assemblage and the pollination of Psychotria nuda (Rubiaceae), a hummingbird-pollinated plant endemic to the Brazilian Atlantic Rainforest. At increasing distance (0–1000 m) from a feeding-station, where hummingbirds have been fed continuously for the past 13 years, we quantified hummingbird abundance, and rates of flower visitation and pollen deposition on P. nuda. We found that hummingbird abundance was unrelated to distance from the feeders beyond ca. 75 m, but increased steeply closer to the feeders; the only exception was the small hummingbird Phaethornis ruber, which remained absent from the feeders. Plants of P. nuda within ca.125 m from the feeders received increasingly more visits, coinciding with the higher hummingbird abundance, whereas visitation rate beyond 125 m showed no distance-related trend. Despite this, pollen deposition was not associated with distance from the feeders. Our findings illustrate that artificial nectar-feeders may locally increase hummingbird abundance, and possibly affect species composition and pollination redundancy, without necessarily having a disruptive effect on pollination services and plants’ reproductive fitness. This may apply not only to hummingbirds, but also to other animal pollinators.

Woody plants host diverse communities of associated organisms, including wood-inhabiting fungi. In this group, host effects on species richness and interaction network structure are not well understood, especially not at large geographical scales. We investigated ecological, historical and evolutionary determinants of fungal species richness and network modularity, that is, subcommunity structure, across woody hosts in Denmark, using a citizen science data set comprising > 80 000 records of > 1000 fungal species on 91 genera of woody plants. Fungal species richness was positively related to host size, wood pH, and the number of species in the host genus, with limited influence of host frequency and host history, that is, time since host establishment in the area. Modularity patterns were unaffected by host history, but largely reflected host phylogeny. Notably, fungal communities differed substantially between angiosperm and gymnosperm hosts. Host traits and evolutionary history appear to be more important than host frequency and recent history in structuring interactions between hosts and wood-inhabiting fungi. High wood acidity appears to act as a stress factor reducing fungal species richness, while large host size, providing increased niche diversity, enhances it. In some fungal groups that are known to interact with live host cells in the establishment phase, host selectivity is common, causing a modular community 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.

Aim: To investigate the role of alien plants in mutualistic plant-hummingbird networks, assessing the importance of species traits, floral abundance and insularity on alien plant integration. Location: Mainland and insular Americas. Methods: We used species-level network indices to assess the role of alien plants in 21 quantitative plant-hummingbird networks where alien plants occur. We then evaluated whether plant traits, including previous adaptations to bird pollination, and insularity predict these network roles. Additionally, for a subset of networks for which floral abundance data were available, we tested whether this relates to network roles. Finally, we tested the association between hummingbird traits and the probability of interaction with alien plants across the networks. Results: Within the 21 networks, we identified 32 alien plant species and 352 native plant species. On average, alien plant species attracted more hummingbird species (i.e. aliens had a higher degree) and had a higher proportion of interactions across their hummingbird visitors than native plants (i.e. aliens had a higher species strength). At the same time, an average alien plant was visited more exclusively by certain hummingbird species (i.e. had a higher level of complementary specialization). Large alien plants and those occurring on islands had more evenly distributed interactions, thereby acting as connectors. Other evaluated plant traits and floral abundance were unimportant predictors of network roles. Short-billed hummingbirds had higher probability of including alien plants in their interactions than long-billed species. Main conclusions: Once incorporated into plant-hummingbird networks, alien plants appear strongly integrated and, thus, may have a large influence on network dynamics. Plant traits and floral abundance were generally poor predictors of how well alien species are integrated. Short-billed hummingbirds, often characterized as functionally generalized pollinators, facilitate the integration of alien plants. Our results show that plant-hummingbird networks are open for invasion. © 2016 John Wiley & Sons Ltd.

Species traits are thought to predict feeding specialization and the vulnerability of a species to extinctions of interaction partners, but the context in which a species evolved and currently inhabits may also matter. Notably, the predictive power of traits may require that traits evolved to fit interaction partners. Furthermore, local abiotic and biotic conditions may be important. On islands, for instance, specialized and vulnerable species are predicted to be found mainly in mountains, whereas species in lowlands should be generalized and less vulnerable. We evaluated these predictions for hummingbirds and their nectar-food plants on Antillean islands. Our results suggest that the rates of hummingbird trait divergence were higher among ancestral mainland forms before the colonization of the Antilles. In correspondence with the limited trait evolution that occurred within the Antilles, local abiotic and biotic conditions—not species traits—correlate with hummingbird resource specialization and the vulnerability of hummingbirds to extinctions of their floral resources. Specifically, hummingbirds were more specialized and vulnerable in conditions with high topographical complexity, high rainfall, low temperatures and high floral resource richness, which characterize the Antillean Mountains. These findings show that resource specialization and species vulnerability to extinctions of interaction partners are highly context-dependent.