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Mutualistic interactions, such as plant–mycorrhizal or plant–pollinator interactions, are widespread in ecological communities and frequently exploited by cheaters, species that profit from interactions without providing benefits in return. Cheating usually negatively affects the fitness of the individuals that are cheated on, but the effects of cheating at the community level remains poorly understood. Here, we describe 2 different kinds of cheating in mutualistic networks and use a generalized Lotka–Volterra model to show that they have very different consequences for the persistence of the community. Conservative cheating, where a species cheats on its mutualistic partners to escape the cost of mutualistic interactions, negatively affects community persistence. In contrast, innovative cheating occurs with species with whom legitimate interactions are not possible, because of a physiological or morphological barrier. Innovative cheating can enhance community persistence under some conditions: when cheaters have few mutualistic partners, cheat at low or intermediate frequency and the cost associated with mutualism is not too high. Under these conditions, the negative effects of cheating on partner persistence are overcompensated at the community level by the positive feedback loops that arise in diverse mutualistic communities. Using an empirical dataset of plant–bird interactions (hummingbirds and flowerpiercers), we found that observed cheating patterns are highly consistent with theoretical cheating patterns found to increase community persistence. This result suggests that the cheating patterns observed in nature could contribute to promote species coexistence in mutualistic communities, instead of necessarily destabilizing them. Mutualistic interactions, such as plant-mycorrhizal or plant-pollinator interactions, are widespread in ecological communities and frequently exploited by cheaters. This combination of theoretical model and empirical data suggests that cheating in pollination could contribute to the stability of plant-pollinator communities [ABSTRACT FROM AUTHOR]

How traits affect speciation is a long-standing question in evolution. We investigate whether speciation rates are affected by the traits themselves or by the rates of their evolution, in hummingbirds, a clade with great variation in speciation rates, morphology and ecological niches. Further, we test two opposing hypotheses, postulating that speciation rates are promoted by trait conservatism or, alternatively, by trait divergence. To address these questions, we analyse morphological (body mass and bill length) and niche traits (temperature and precipitation position and breadth, and mid-elevation), using a variety of methods to estimate speciation rates and correlate them with traits and their evolutionary rates. When it comes to the traits, we find faster speciation in smaller hummingbirds with shorter bills, living at higher elevations and experiencing greater temperature ranges. As for the trait evolutionary rates, we find that speciation increases with rates of divergence in the niche traits, but not in the morphological traits. Together, these results reveal the interplay of mechanisms through which different traits and their evolutionary rates (conservatism or divergence) influence the origination of hummingbird diversity. [ABSTRACT FROM AUTHOR]

Niche partitioning is an important mechanism that allows species to coexist. Within mutualistic interaction networks, diel niche partitioning, i.e., partitioning of resources throughout the day, has been neglected. We explored diel niche partitioning of a plant-hummingbird network in the Brazilian Atlantic forest for nine months. To evaluate diel patterns of hummingbird visits and nectar production, we used time-lapse cameras on focal flowers and repeated nectar volume and concentration measures, respectively. Additionally, we measured flower abundance around focal flowers and flower morphological traits. We did not observe diel partitioning for either hummingbirds or plants. Instead, hummingbirds appeared to specialize in different plant species, consistent with trophic niche partitioning, potentially resulting from competition. In contrast, plant species that co-flowered and shared hummingbird visits produced nectar during similar times, consistent with facilitation. Our focus on the fine-scale temporal pattern revealed that plants and hummingbirds appear to have different strategies for promoting co-existence. [ABSTRACT FROM AUTHOR]

On-going land-use change has profound impacts on biodiversity by filtering species that cannot survive in disturbed landscapes and potentially altering biotic interactions. In particular, how land-use change reshapes biotic interactions remains an open question. Here, we used selectivity experiments with nectar feeders in natural and converted forests to test the direct and indirect effects of land-use change on resource competition in Andean hummingbirds along an elevational gradient. Selectivity was defined as the time hummingbirds spent at high resource feeders when feeders with both low and high resource values were offered in the presence of other hummingbird species. Selectivity approximates the outcome of interspecific competition (i.e., the resource intake across competing species); in the absence of competition, birds should exhibit higher selectivity. We evaluated the indirect effect of forest conversion on selectivity, as mediated by morphological dissimilarity and flower resource abundance, using structural equation models. We found that forest conversion influenced selectivity at low and mid-elevations, but the influence of morphological dissimilarity and resource availability on selectivity varied between these elevations. At mid-elevation, selectivity was more influenced by the presence of morphologically similar competitors than by resource abundance while at low-elevation resource abundance was a more important predictor of selectivity. Our results suggest that selectivity is influenced by forest conversion, but that the drivers of these changes vary across elevation, highlighting the importance of considering context-dependent variation in the composition of resources and competitors when studying competition. [ABSTRACT FROM AUTHOR]

Torpor is an important energy saving strategy in some small birds, but it has rarely been studied in natural field conditions. We compared torpor use across 43 wild‐caught individuals of eight hummingbird species across sites with different natural temperature regimes. Most laboratory studies focus on the relationship between metabolic rate and temperature, but our aim was to evaluate what environmental factors most influence hummingbird nighttime energy management under natural conditions. We found that the probability of an individual entering torpor was weakly correlated with mass but unrelated to nighttime temperature and that hummingbirds at both warm, tropical and cooler, temperate sites used torpor. Energy savings in torpor were maximized as ambient temperatures approached a species' minimum body temperature, consistent with laboratory studies; energy savings ranged between 65 and 92% of energy per hour in torpor compared to normothermy. However, regardless of the degree of energy savings in torpor, variation in total nighttime energy expenditure was most significantly influenced by torpor bout duration. Lab studies largely assess the effect of temperature on torpor use, but our findings indicate that other environmental conditions are more important in determining hummingbirds' total nighttime energy expenditure under natural temperature cycles. Our results show that a small endotherm's nighttime energy management in its natural habitat is more affected by torpor bout duration, which is linked to photoperiod, than by temperature. This result suggests that in their natural environments hummingbirds are able to save energy in torpor across a range of nighttime temperatures, indicating that they may have sufficient physiological flexibility to tolerate climatic variation. [ABSTRACT FROM AUTHOR]

Within‐clade allometric relationships represent standard laws of scaling between energy and size, and their outliers provide new avenues for physiological and ecological research. According to the metabolic‐level boundaries hypothesis, metabolic rates as a function of mass are expected to scale closer to 0.67 when driven by surface‐related processes (e.g. heat or water flux), while volume‐related processes (e.g. activity) generate slopes closer to one.In birds, daily energy expenditure (DEE) scales with body mass (M) in the relationship logDEE=2.35+0.68×logM, consistent with surface‐level processes driving the relationship. However, taxon‐specific patterns differ from the scaling slope of all birds.Hummingbirds have the highest mass‐specific metabolic rates among all vertebrates. Previous studies on a few hummingbird species, without accounting for the phylogeny, estimated that the DEE–body mass relationship for hummingbirds was logDEE=1.72+1.21×logM. In Contrast to the theoretical expectations, this slope >1 indicates that larger hummingbirds are less metabolically efficient than smaller hummingbirds.We collected DEE and mass data for 12 hummingbird species, which, combined with published data, represented 17 hummingbird species in eight of nine hummingbird clades over a sixfold size range of body size (2.7–17.5 g).After accounting for phylogenetic relatedness, we found DEE scales with body mass as logDEE=2.04+0.95×logM. This slope of 0.95 is lower than previously estimated for hummingbirds, but much higher than the slope for all birds (0.68). The high slopes of torpor, hovering and flight potentially explain the high interspecific DEE slope for hummingbirds compared to other endotherms. [ABSTRACT FROM AUTHOR]

A key component of individual fitness is the ability to manage energy stores in response to variable resource availability, but because directly measuring energy budgets is difficult, daily energy management is rarely measured.Hummingbirds' energy management is relatively simple to model compared to other endotherms because they have high mass‐specific metabolic rates and store little fat.We determined which aspects of the hummingbird daily energy budget (i.e. thermoregulation, daytime activity costs, night‐time costs) change at the individual level in response to environmental variation.We found that daily energy expenditure varied threefold in two populations of broad‐billed hummingbirds (Cynanthus latirostris).Our model indicated the energy budget was distributed in the following proportions: daytime activity, 59% (range 22%–84%); thermoregulation, 23% (11%–32%); basal metabolism, 7% (3%–16%); and night‐time energy, 17% (6%–37%). Activity costs were higher at the hotter, homogeneous site and during the early‐wet season at both sites.Increased daily energy expenditure was related to decreased nectar availability and not significantly related to temperature or bird mass. With climate change, the indirect energetic costs of shifting resources could have greater impacts on endotherm energy budgets than direct costs such as thermoregulation. Increased foraging and activity costs could decrease the energy available to birds for somatic repair and reproduction, potentially causing differential fitness across seasons and sites. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

High-elevation organisms experience shared environmental challenges that include low oxygen availability, cold temperatures, and intense ultraviolet radiation. Consequently, repeated evolution of the same genetic mechanisms may occur across high-elevation taxa. To test this prediction, we investigated the extent to which the same biochemical pathways, genes, or sites were subject to parallel molecular evolution for 12 Andean hummingbird species (family: Trochilidae) representing several independent transitions to high elevation across the phylogeny. Across high-elevation species, we discovered parallel evolution for several pathways and genes with evidence of positive selection. In particular, positively selected genes were frequently part of cellular respiration, metabolism, or cell death pathways. To further examine the role of elevation in our analyses, we compared results for low- and high-elevation species and tested different thresholds for defining elevation categories. In analyses with different elevation thresholds, positively selected genes reflected similar functions and pathways, even though there were almost no specific genes in common. For example, EPAS1 (HIF2α), which has been implicated in high-elevation adaptation in other vertebrates, shows a signature of positive selection when high-elevation is defined broadly (>1,500 m), but not when defined narrowly (>2,500 m). Although a few biochemical pathways and genes change predictably as part of hummingbird adaptation to high-elevation conditions, independent lineages have rarely adapted via the same substitutions. [ABSTRACT FROM AUTHOR]

Abstract: Species interactions are fundamental to community dynamics and ecosystem processes. Despite significant progress in describing species interactions, we lack the ability to predict changes in interactions across space and time. We outline a Bayesian approach to separate the probability of species co‐occurrence, interaction and detectability in influencing interaction betadiversity. We use a multi‐year hummingbird–plant time series, divided into training and testing data, to show that including models of detectability and occurrence improves forecasts of mutualistic interactions. We then extend our model to explore interaction betadiversity across two distinct seasons. Despite differences in the observed interactions among seasons, there was no significant change in hummingbird occurrence or interaction frequency between hummingbirds and plants. These results highlight the challenge of inferring the causes of interaction betadiversity when interaction detectability is low. Finally, we highlight potential applications of our model for integrating observations of local interactions with biogeographic and evolutionary histories of co‐occurring species. These advances will provide new insight into the mechanisms that drive variation in patterns of biodiversity. [ABSTRACT FROM AUTHOR]

Land use change modifies the environment at multiple spatial scales, and is a main driver of species declines and deterioration of ecosystem services. However, most of the research on the effects of land use change has focused on taxonomic diversity, while functional diversity, an important predictor of ecosystem services, is often neglected. We explored how local and landscape scale characteristics influence functional and taxonomic diversity of hummingbirds in the Andes Mountains in southern Ecuador. Data was collected in six landscapes along a land use gradient, from an almost intact landscape to one dominated by cattle pastures. We used point counts to sample hummingbirds from 2011 to 2012 to assessed how local factors (i.e., vegetation structure, flowering plants richness, nectar availability) and landscape factors (i.e., landscape heterogeneity, native vegetation cover) influenced taxonomic and functional diversity. Then, we analyzed environment - trait relationships (RLQ test) to explore how different hummingbird functional traits influenced species responses to these factors. Taxonomic and functional diversity of hummingbirds were positively associated with landscape heterogeneity but only functional diversity was positively related to native vegetation coverage. We found a weak response of taxonomic and functional diversity to land use change at the local scale. Environment-trait associations showed that body mass of hummingbirds likely influenced species sensitivity to land use change. In conclusion, landscape heterogeneity created by land use change can positively influence hummingbird taxonomic and functional diversity; however, a reduction of native vegetation cover could decrease functional diversity. Given that functional diversity can mediate ecosystem services, the conservation of native vegetation cover could play a key role in the maintenance of hummingbird pollination services in the tropical Andes. Moreover, there are particular functional traits, such as body mass, that increase a species sensitivity to land use change. [ABSTRACT FROM AUTHOR]

The composition of ecological assemblages depends on a variety of factors including environmental filtering, biotic interactions and dispersal limitation. By evaluating the phylogenetic pattern of assemblages, we gain insight into the relative contribution of these mechanisms to generating observed assemblages. We address some limitations in the field of community phylogenetics by using simulations, biologically relevant null models, and cost distance analysis to evaluate simultaneous mechanisms leading to observed patterns of co-occurrence. Building from past studies of phylogenetic community structure, we applied our approach to hummingbird assemblages in the Northern Andes. We compared the relationship between relatedness and co-occurrence among predicted assemblages, based on estimates of suitable habitat and dispersal limitation, and observed assemblages. Hummingbird co-occurrence peaked at intermediate relatedness and decreased when a closely-related species was present. This result was most similar to simulations that included simultaneous effects of phylogenetic conservatism and repulsion. In addition, we found older sister taxa were only weakly more separated by geographic barriers, suggesting that time since dispersal is unlikely to be the sole factor influencing co-occurrence of closely related species. Our analysis highlights the role of multiple mechanisms acting simultaneously, and provides a hypothesis for the potential importance of competition at regional scales. [ABSTRACT FROM AUTHOR]

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Aim Under climate change, it is likely that as species reshuffle based on their environmental tolerances, novel assemblages will form and some current assemblages will disappear. It is important for future monitoring and conservation that we understand where these novel and disappearing assemblages occur and how they differ among dimensions (taxonomic, phylogenetic and functional) of diversity. Here we investigate the geographical and environmental patterns of novel and disappearing assemblages; whether these patterns hold across dimensions of diversity; and how these assemblages are characterized in trait space. Location Ecuador. Methods We used ensemble species distribution modelling to estimate the distributions of 151 hummingbird species into the projected climate for 2070. Using standard beta diversity measures, we identified novel and disappearing taxonomic, phylogenetic and functional assemblages. Results We found that novel and disappearing hummingbird assemblages are likely under climate change, particularly in extreme environments and with novel assemblages replacing disappearing assemblages. Although the patterns of novel and disappearing assemblages were similar among dimensions of diversity, we found that there were fewest novel and disappearing functional assemblages. The future assemblages were characterized by an increase in functional space, which is counter to typical predictions of trait homogenization under climate change. Main conclusions Novel and disappearing assemblages are likely to pose management challenges for future conservation. Here we present an approach to identify such assemblages. By considering the geographic and environmental context of novel and disappearing assemblages for different dimensions of diversity, we can start to identify the mechanisms behind these patterns. [ABSTRACT FROM AUTHOR]

By specialising on specific resources, species evolve advantageous morphologies to increase the efficiency of nutrient acquisition. However, many specialists face variation in resource availability and composition. Whether specialists respond to these changes depends on the composition of the resource pulses, the cost of foraging on poorly matched resources, and the strength of interspecific competition. We studied hummingbird bill and plant corolla matching during seasonal variation in flower availability and morphology. Using a hierarchical Bayesian model, we accounted for the detectability and spatial overlap of hummingbird-plant interactions. We found that despite seasonal pulses of flowers with short-corollas, hummingbirds consistently foraged on well-matched flowers, leading to low niche overlap. This behaviour suggests that the costs of searching for rare and more specialised resources are lower than the benefit of switching to super-abundant resources. Our results highlight the trade-off between foraging efficiency and interspecific competition, and underline niche partitioning in maintaining tropical diversity. [ABSTRACT FROM AUTHOR]

Specialization of species in interaction networks influences network stability and ecosystem functioning. Spatial and temporal variation in resource availability may provide insight into how ecological factors, such as resource abundance, and evolutionary factors, such as phylogenetically conserved morphological traits, influence specialization within mutualistic networks. We used independent measures of hummingbird abundance and resources (nectar), information on hummingbird traits and plant-hummingbird interactions to examine how resource availability and species' morphology influence the specialization of hummingbirds in three habitat types (forest, shrubs, cattle ranch) sampled over 10 sessions across two years in the southern Andes of Ecuador. Specialization of hummingbird species in the networks was measured by three indices: d' (related to niche partitioning), generality (related to niche width) and PSI (related to pollination services). Specialization indices d', generality and PSI of hummingbird species were influenced by resource availability. All indices indicated that specialization of hummingbirds increased when the availability of resources decreased. Variation in d' was also explained by an interaction between resource availability and bill length; hummingbirds with a long bill switched from being more specialized than other species when resource availability was low to being similarly specialized when availability was high. Overall, our results highlight the importance of ecological and evolutionary factors determining the specialization of species in interaction networks. We demonstrate in particular that ecological gradients in resource availability cause substantial changes in consumers' foraging behavior contingent on their morphology. Changes in pollinator specialization along resource gradients can have impacts on ecosystem functions, such as pollination by animals. [ABSTRACT FROM AUTHOR]

A challenge in community ecology is connecting biogeographic patterns with local scale observations. In Neotropical hummingbirds, closely related species often co-occur less frequently than expected (overdispersion) when compared to a regional species pool. While this pattern has been attributed to interspecific competition, it is important to connect these findings with local scale mechanisms of coexistence. We measured the importance of the presence of competitors and the availability of resources on selectivity at experimental feeders for Andean hummingbirds along a wide elevation gradient. Selectivity was measured as the time a bird fed at a feeder with a high sucrose concentration when presented with feeders of both low and high sucrose concentrations. Resource selection was measured using time-lapse cameras to identity which floral resources were used by each hummingbird species. We found that the increased abundance of preferred resources surrounding the feeder best explained increased species selectivity, and that related hummingbirds with similar morphology chose similar floral resources. We did not find strong support for direct agonism based on differences in body size or phylogenetic relatedness in predicting selectivity. These results suggest closely related hummingbird species have overlapping resource niches, and that the intensity of interspecific competition is related to the abundance of those preferred resources. If these competitive interactions have negative demographic effects, our results could help explain the pattern of phylogenetic overdispersion observed at regional scales. [ABSTRACT FROM AUTHOR]

A persistent challenge in ecology is to tease apart the influence of multiple processes acting simultaneously and interacting in complex ways to shape the structure of species assemblages. We implement a heuristic approach that relies on explicitly defining species pools and permits assessment of the relative influence of themain processes thought to shape assemblage structure: environmental filtering, dispersal limitations, and biotic interactions. We illustrate our approach using data on the assemblage composition and geographic distribution of hummingbirds, a comprehensive phylogeny and morphological traits. The implementation of several process-based species pool definitions in null models suggests that temperature—but not precipitation or dispersal limitation—acts as the main regional filter of assemblage structure. Incorporating this environmental filter directly into the definition of assemblage-specific species pools revealed an otherwise hidden pattern of phylogenetic evenness, indicating that biotic interactions might further influence hummingbird assemblage structure. Such hidden patterns of assemblage structure call for a reexamination of a multitude of phylogenetic- and trait-based studies that did not explicitly consider potentially important processes in their definition of the species pool. Our heuristic approach provides a transparent way to explore patterns and refine interpretations of the underlying causes of assemblage structure. [ABSTRACT FROM AUTHOR]

Aim To investigate the association between hummingbird-plant network structure and species richness, phylogenetic signal on species' interaction pattern, insularity and historical and current climate. Location Fifty-four communities along a c. 10,000 km latitudinal gradient across the Americas (39° N-32° S), ranging from sea level to c. 3700 m a.s.l., located on the mainland and on islands and covering a wide range of climate regimes. Methods We measured the level of specialization and modularity in mutualistic plant-hummingbird interaction networks. Using an ordinary least squares multimodel approach, we examined the influence of species richness, phylogenetic signal, insularity and current and historical climate conditions on network structure (null-model-corrected specialization and modularity). Results Phylogenetically related species, especially plants, showed a tendency to interact with a similar array of mutualistic partners. The spatial variation in network structure exhibited a constant association with species phylogeny ( R2 = 0.18-0.19); however, network structure showed the strongest association with species richness and environmental factors ( R2 = 0.20-0.44 and R2 = 0.32-0.45, respectively). Specifically, higher levels of specialization and modularity were associated with species-rich communities and communities in which closely related hummingbirds visited distinct sets of flowering species. On the mainland, specialization was also associated with warmer temperatures and greater historical temperature stability. Main conclusions Our results confirm the results of previous macroecological studies of interaction networks which have highlighted the importance of species richness and the environment in determining network structure. Additionally, for the first time, we report an association between network structure and species phylogenetic signal at a macroecological scale, indicating that high specialization and modularity are associated with high interspecific competition among closely related hummingbirds, subdividing the floral niche. This suggests a tighter co-evolutionary association between hummingbirds and their plants than in previously studied plant-bird mutualistic systems. [ABSTRACT FROM AUTHOR]

Patterns of phylogenetic structure of assemblages are increasingly used to gain insight into the ecological and evolutionary processes involved in the assembly of co-occurring species. Metrics of phylogenetic structure can be sensitive to scaling issues and data availability. Here we empirically assess the sensitivity of four metrics of phylogenetic structure of assemblages to changes in (i) the source of data, (ii) the spatial grain at which assemblages are defined, and (iii) the definition of species pools using hummingbird (Trochilidae) assemblages along an elevational gradient in Colombia. We also discuss some of the implications in terms of the potential mechanisms driving these patterns. To explore how source of data influence phylogenetic structure we defined assemblages using three sources of data: field inventories, museum specimens, and range maps. Assemblages were defined at two spatial grains: coarse-grained (elevational bands of 800-m width) and fine-grained (1-km2 plots). We used three different species pools: all species contained in assemblages, all species within half-degree quadrats, and all species either above or below 2000 m elevation. Metrics considering phylogenetic relationships among all species within assemblages showed phylogenetic clustering at high elevations and phylogenetic evenness in the lowlands, whereas those metrics considering only the closest co-occurring relatives showed the opposite trend. This result suggests that using multiple metrics of phylogenetic structure should provide greater insight into the mechanisms shaping assemblage structure. The source and spatial grain of data had important influences on estimates of both richness and phylogenetic structure. Metrics considering the co-occurrence of close relatives were particularly sensitive to changes in the spatial grain. Assemblages based on range maps included more species and showed less phylogenetic structure than assemblages based on museum or field inventories. Coarse-grained assemblages included more distantly related species and thus showed a more even phylogenetic structure than fine-grained assemblages. Our results emphasize the importance of carefully selecting the scale, source of data and metric used in analysis of the phylogenetic structure of assemblages. [ABSTRACT FROM AUTHOR]