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Indirect interactions are pivotal in the evolution of interacting species and the assembly of populations and communities. Nevertheless, despite recently being investigated in plant-animal mutualism at the community level, indirect interactions have not been studied in resource-mediated mutualisms involving plant individuals that share different animal species as partners within a population (i.e., individual-based networks). Here, we analyzed an individual-based ant-plant network to evaluate how resource properties affect indirect interaction patterns and how changes in indirect links leave imprints in the network across multiple levels of network organization. Using complementary analytical approaches, we described the patterns of indirect interactions at the micro-, meso-, and macro-scale. We predicted that plants offering intermediate levels of nectar quantity and quality interact with more diverse ant assemblages. The increased number of ant species would cause a higher potential for indirect interactions in all scales evaluated. We found that nectar properties modified patterns of indirect interactions of plant individuals that share mutualistic partners, leaving imprints across different network scales. To our knowledge, this is the first study tracking indirect interactions in multiple scales within an individual-based network. We show that functional traits of interacting species, such as nectar properties, may lead to changes in indirect interactions, which could be tracked across different levels of the network organization evaluated.

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ABSTRACT Ecological interactions are diverse, variable across space and time and not always well understood. The use of interaction network analysis has become a tool that promotes a deeper understanding on ecological and evolutionary processes. The interaction between insects and fungi is an interesting research model, helping to understand colonization dynamics and species specialization in spatially aggregated and ephemeral resources. Here, we describe the interactions between Drosophilidae species and the fungal basidiocarps in a subtropical forest in Brazil. Flies were collected when were visiting basidiocarps and then the basidiocarps themselves were also collected to obtain the emerging flies whose larvae fed on the fungi. We observed 31 species of drosophilids interacting with basidiocarps of 23 fungi species. An ecological network analysis was performed for the drosophilids breeding on basidiocarps and for those visiting them as adults. We found a specialized breeding network, with stronger interactions involving Hirtodrosophila and Auricularia and Zygothrica bilineata and a Marasmius species. Our results indicate the generalist habit of most Zygothrica species. The visitation network was highly specialized. Despite being well represented in the sampling, most Zygothrica species did not emerge from any fungal species. This study advances the knowledge on patterns of Drosophilid-fungi interactions and provides insights into their drivers.

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Research Highlight: Leimberger, K.G., Hadley, A.S., & Betts, M.G. (2023). Plant-hummingbird pollination networks exhibit minimal rewiring after experimental removal of a locally abundant plant species. Journal of Animal Ecology, https://doi.org/10.1111/1365-2656.13935. In this paper, Leimberger, Hadley and Betts (2023) explore the effects of removing a locally abundant plant species on plant-hummingbird pollination networks. They experimentally prevented access of hummingbirds to flowers of Heliconia tortuosa and assessed subsequent changes in the interactions between plants and hummingbirds. Their main hypothesis postulated that the loss of a highly connected species would lead to interaction rewiring and niche expansions by hummingbirds, decreasing individual, species and network specialization. However, they found that the overall structure of the plant-hummingbird networks remains mostly unaltered, with limited rewiring and minimal changes in specialization. The main contributions of this study can be summarized as (i) it adds to a limited number of manipulative studies on the capacity of species to rewire their interactions following the loss of partners, and importantly, it is the first study from the tropics and with vertebrate pollinators, for which experimental studies at appropriate scales is intrinsically more challenging; and (ii) innovates by evaluating change in specialization for the individual level, carried out through pollen sampling on the body of hummingbirds. The limited change in species interactions highlights that network stability through interaction rewiring may have been overestimated in previous studies, calling for further manipulative studies in the field. At the same time, it also indicated that even the loss of a highly abundant plant species has an overall small effect on network structure. Thus, this study contributes timely findings regarding the capacity of ecological communities to respond to species extinctions.

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Understanding the mechanisms that shape the architecture, diversity, and adaptations of genomes and their ecological and genetic interfaces is of utmost importance to understand biological evolution. Transposable elements (TEs) play an important role in genome evolution, due to their ability to transpose within and between genomes, providing sites of nonallelic recombination. Here we investigate patterns and processes of TE-driven genome evolution associated with niche diversification. Specifically, we compared TE content, TE landscapes, and frequency of horizontal transposon transfers (HTTs) across genomes of flower-breeding Drosophila (FBD) with different levels of specialization on flowers. Further, we investigated whether niche breadth and ecological and geographical overlaps are associated with a potential for HTT rates. Landscape analysis evidenced a general phylogenetic pattern, in which species of the D. bromeliae group presented L-shaped curves, indicating recent transposition bursts, whereas D. lutzii showed a bimodal pattern. The great frequency of highly similar sequences recovered for all FBD suggests that these species probably experienced similar ecological pressures and evolutionary histories that contributed to the diversification of their mobilomes. Likewise, the richness of TEs superfamilies also appears to be associated with ecological traits. Furthermore, the two more widespread species, the specialist D. incompta and the generalist D. lutzii, presented the highest frequency of HTT events. Our analyses also revealed that HTT opportunities are positively influenced by abiotic niche overlap but are not associated with phylogenetic relationships or niche breadth. This suggests the existence of intermediate vectors promoting HTTs between species that do not necessarily present overlapping biotic niches.

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As human-caused extinctions and invasions accumulate across the planet, understanding the processes governing ecological functions mediated by species interactions, and anticipating the effect of species loss on such functions become increasingly urgent. In seed dispersal networks, the mechanisms that influence interaction frequencies may also influence the capacity of a species to switch to alternative partners (rewiring), influencing network robustness. Studying seed dispersal interactions in novel ecosystems on O'ahu island, Hawai'i, we test whether the same mechanisms defining interaction frequencies can regulate rewiring and increase network robustness to simulated species extinctions. We found that spatial and temporal overlaps were the primary mechanisms underlying interaction frequencies, and the loss of the more connected species affected networks to a greater extent. Further, rewiring increased network robustness, and morphological matching and spatial and temporal overlaps between partners were more influential on network robustness than species abundances. We argue that to achieve self-sustaining ecosystems, restoration initiatives can consider optimal morphological matching and spatial and temporal overlaps between consumers and resources to maximize chances of native plant dispersal. Specifically, restoration initiatives may benefit from replacing invasive species with native species possessing characteristics that promote frequent interactions and increase the probability of rewiring (such as long fruiting periods, small seeds and broad distributions).

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While network analyses have stimulated a renewed interest in understanding patterns and drivers of specialization within communities, few studies have explored specialization within populations. Thus, in plant populations, causes and consequences of individual variation in their interactions with mutualistic animals remain poorly understood. Studying a Brazilian pepper (Schinus terebinthifolia) population, we measured the extent of individual variation in interactions with seed dispersers and tested whether connectivity (number of seed dispersers) and specialization (exclusiveness of partners) are associated with phenotypic and phenological traits of individuals and their spatial context. We found that: (i) individuals varied broadly in their connectivity and specialization on seed dispersers; (ii) phenotypic traits and spatial context matter more than fruiting duration in determining how many and how exclusive are seed dispersers of an individual; (iii) the individual-based network was nested and indicated that the less connected individuals were shorter, occurred in neighborhoods with fewer fruits, and tended to interact with a subset of the partners of more generalist individuals which, in turn, were taller and inserted in higher fruit density neighborhoods; (iv) modularity indicated the existence of subsets of individuals that interacted disproportionately with distinct groups of partners, which may occur due to differences in bird habitat use across the landscape. Our study underlines a remarkable interindividual variation that is overlooked when interactions are compiled to describe species-level interactions. Traits and spatial contexts that define variation among individuals may have important implications not only for fitness but also for sampling and description of interactions at species level.

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Flowering plant species and their nectar-feeding vertebrates exemplify some of the most remarkable biotic interactions in the Neotropics. In the Brazilian Atlantic Forest, several species of birds (especially hummingbirds), bats, and non-flying mammals, as well as one lizard feed on nectar, often act as pollinators and contribute to seed output of flowering plants. We present a dataset containing information on flowering plants visited by nectar-feeding vertebrates and sampled at 166 localities in the Brazilian Atlantic Forest. This dataset provides information on 1902 unique interactions among 515 species of flowering plants and 129 species of potential vertebrate pollinators and the patterns of species diversity across latitudes. All plant-vertebrate interactions compiled were recorded through direct observations of visits, and no inferences of pollinators based on floral syndromes were included. We also provide information on the most common plant traits used to understand the interactions between flowers and nectar-feeding vertebrates: plant growth form, corolla length, rate of nectar production per hour in bagged flowers, nectar concentration, flower color and shape, time of anthesis, presence or absence of perceptible fragrance by human, and flowering phenology as well as the plant's threat status by International Union for Conservation of Nature (IUCN) classification. For the vertebrates, status of threat by IUCN classification, body mass, bill or rostrum size are provided. Information on the frequency of visits and pollen deposition on the vertebrate's body is provided from the original source when available. The highest number of unique interactions is recorded for birds (1771) followed by bats (110). For plants, Bromeliaceae contains the highest number of unique interactions (606), followed by Fabaceae (242) and Gesneriaceae (104). It is evident that there was geographical bias of the studies throughout the southeast of the Brazilian Atlantic Forest and that most effort was directed to flower-hummingbird interactions. However, it reflects a worldwide tendency of more plants interacting with birds compared with other vertebrate species. The lack of similar protocols among studies to collect basic data limits the comparisons among areas and generalizations. Nevertheless, this dataset represents a notable effort to organize and highlight the importance of vertebrate pollinators in this hotspot of biodiversity on Earth and represents the data currently available. No copyright or proprietary restrictions are associated with the use of this data set. Please cite this data paper when the data are used in publications or scientific events.

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Fire is a frequent disturbance in most grasslands around the world, being key for the structure and dynamics of the biodiversity in such ecosystems. While grassland species may be resilient, little is known on how plant-pollinator networks reassemble after fire. Here, we investigate the structure and dynamics of plant-pollinator networks and the variation in species roles over a 2-year post-fire chronosequence on grassland communities in Southern Brazil. We found that both network specialization and modularity were similar over the chronosequence of time-since-fire, but in freshly burnt areas, there were more species acting as network hubs. Species roles exhibited high variation, with plant and pollinator species shifting roles along the post-disturbance chronosequence. Interaction dissimilarity was remarkably high in networks irrespective of times-since-fire. Interaction dissimilarity was associated more with rewiring than with species turnover, indicating that grassland plant and pollinator species are highly capable of switching partners. Time-since-fire had little influence on network structure but influenced the identity and diversity of pollinators playing key roles in the networks. These findings suggest that pollination networks in naturally fire-prone ecosystems are highly dynamic and resilient to fire with both plants and pollinators being highly capable of adjusting their interactions and network structure after disturbance.

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Patterns of specialization and the structure of interactions between bats and ectoparasitic flies have been studied mostly on non-urban environments and at local scales. Thus, how anthropogenic disturbances influence species interactions and network structure in this system remain poorly understood. Here, we investigated patterns of interaction between Phyllostomidae bats and ectoparasitic Streblidae flies, and variations in network specialization and structure across Cerrado patches within urbanized landscapes in Brazil and between local and regional scales. We found high similarity in the richness and composition of bat and fly species across communities, associated with low turnover of interactions between networks. The high specialization of bat-streblid interactions resulted in little connected and modular networks, with the emergence of modules containing subsets of species that interact exclusively or primarily with each other. Such similarities in species and interaction composition and network structure across communities and scales suggest that bat-fly interactions within Cerrado patches are little affected by the degree of human modification in the surrounding matrix. This remarkable consistency is likely promoted by specific behaviors, the tolerance of Phyllostomidae bats to surrounding urbanized landscapes as well as by the specificity of the streblid-bat interactions shaped over evolutionary time.

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In semi-arid environments, the marked contrast in temperature and precipitation over the year strongly shapes ecological communities. The composition of species and their ecological interactions within a community may vary greatly over time. Although intra-annual variations are often studied, empirical information on how plant-bird relationships are structured within and among years, and how their drivers may change over time are still limited. In this study, we analyzed the temporal dynamics of the structure of plant-hummingbird interaction networks by evaluating changes in species richness, diversity of interactions, modularity, network specialization, nestedness, and ß-diversity of interactions throughout four years in a Mexican xeric shrubland landscape. We also evaluated if the relative importance of abundance, phenology, morphology, and nectar sugar content consistently explains the frequency of pairwise interactions between plants and hummingbirds across different years. We found that species richness, diversity of interactions, nestedness, and network specialization did vary within and among years. We also observed that the ß-diversity of interactions was high among years and was mostly associated with species turnover (i.e., changes in species composition), with a minor contribution of interaction rewiring (i.e., shifting partner species at different times). Finally, the temporal co-occurrence of hummingbird and plant species among months was the best predictor of the frequency of pairwise interactions, and this pattern was consistent within and among years. Our study underscores the importance of considering the temporal scale to understand how changes in species phenologies, and the resulting temporal co-occurrences influence the structure of interaction networks.

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