Pollinators contribute to the maintenance of flowering plant diversity.

Mechanisms that favour rare species are key to the maintenance of diverse communities1-3. One of the most critical tasks for conservation of flowering plant biodiversity is to understand how plant-pollinator interactions contribute to the maintenance of rare species4-7. Here we show that niche partitioning in pollinator use and asymmetric facilitation confer fitness advantage of rarer species in a biodiversity hotspot using phylogenetic structural equation modelling that integrates plant-pollinator and interspecific pollen transfer networks with floral functional traits. Co-flowering species filtered pollinators via floral traits, and rarer species showed greater pollinator specialization leading to higher pollination-mediated male and female fitness than more abundant species. When plants shared pollinator resources, asymmetric facilitation via pollen transport dynamics benefitted the rarer species at the cost of more abundant species, serving as an alternative diversity-promoting mechanism. Our results emphasize the importance of community-wide plant-pollinator interactions that affect reproduction for biodiversity maintenance.

Polyploidy: an evolutionary and ecological force in stressful times.

Polyploidy has been hypothesized to be both an evolutionary dead-end and a source for evolutionary innovation and species diversification. Although polyploid organisms, especially plants, abound, the apparent nonrandom long-term establishment of genome duplications suggests a link with environmental conditions. Whole-genome duplications seem to correlate with periods of extinction or global change, while polyploids often thrive in harsh or disturbed environments. Evidence is also accumulating that biotic interactions, for instance, with pathogens or mutualists, affect polyploids differently than nonpolyploids. Here, we review recent findings and insights on the effect of both abiotic and biotic stress on polyploids versus nonpolyploids and propose that stress response in general is an important and even determining factor in the establishment and success of polyploidy.

Neopolyploidy-induced changes in giant duckweed (Spirodela polyrhiza) alter herbivore preference and performance and plant population performance.

PREMISE: Polyploidy is a widespread mutational process in angiosperms that may alter population performance of not only plants but also their interacting species. Yet, knowledge of whether polyploidy affects plant-herbivore dynamics is scarce. Here, we tested whether aphid herbivores exhibit preference for diploid or neopolyploid plants, whether polyploidy impacts plant and herbivore performance, and whether these interactions depend on the plant genetic background. METHODS: Using independently synthesized neotetraploid strains paired with their diploid progenitors of greater duckweed (Spirodela polyrhiza), we evaluated the effect of neopolyploidy on duckweed's interaction with the water-lily aphid (Rhopalosiphum nymphaeae). Using paired-choice experiments, we evaluated feeding preference of the herbivore. We then evaluated the consequences of polyploidy on aphid and plant performance by measuring population growth over multiple generations. RESULTS: Aphids preferred neopolyploids when plants were provided at equal abundances but not at equal surface areas, suggesting the role of plant population surface area in driving this preference. Additionally, neopolyploidy increased aphid population performance, but this result was dependent on the plant's genetic lineage. Lastly, the impact of herbivory on neopolyploid vs. diploid duckweed varied greatly with genetic lineage, where neopolyploids appeared to be variably tolerant compared to diploids, sometimes mirroring the effect on herbivore performance. CONCLUSIONS: By experimentally testing the impacts of polyploidy on trophic species interactions, we showed that polyploidization can impact the preference and performance of herbivores on their plant hosts. These results have significant implications for the establishment and persistence of plants and herbivores in the face of plant polyploidy.

Neopolyploidy has variable effects on the diversity and composition of the wild strawberry microbiome.

PREMISE: Whole-genome duplication (neopolyploidy) can instantly differentiate the phenotype of neopolyploids from their diploid progenitors. These phenotypic shifts in organs such as roots and leaves could also differentiate the way neopolyploids interact with microbial species. While some studies have addressed how specific microbial interactions are affected by neopolyploidy, we lack an understanding of how genome duplication affects the diversity and composition of microbial communities. METHODS: We performed a common garden experiment with multiple clones of artificially synthesized autotetraploids and their ancestral diploids, derived from 13 genotypes of wild strawberry, Fragaria vesca. We sequenced epiphytic bacteria and fungi from roots and leaves and characterized microbial communities and leaf functional traits. RESULTS: Autotetraploidy had no effect on bacterial alpha diversity of either organ, but it did have a genotype-dependent effect on the diversity of fungi on leaves. In contrast, autotetraploidy restructured the community composition of leaf bacteria and had a genotype-dependent effect on fungal community composition in both organs. The most differentially abundant bacterial taxon on leaves belonged to the Sphingomonas, while a member of the Trichoderma was the most differentially abundant fungal taxon on roots. Ploidy-induced change in leaf size was strongly correlated with a change in bacterial but not fungal leaf communities. CONCLUSIONS: Genome duplication can immediately alter aspects of the plant microbiome, but this effect varies by host genotype and bacterial and fungal community. Expanding these studies to wild settings where plants are exposed continuously to microbes are needed to confirm the patterns observed here.

Polyploidy impacts population growth and competition with diploids: multigenerational experiments reveal key life-history trade-offs.

Ecological theory predicts that early generation polyploids ('neopolyploids') should quickly go extinct owing to the disadvantages of rarity and competition with their diploid progenitors. However, polyploids persist in natural habitats globally. This paradox has been addressed theoretically by recognizing that reproductive assurance of neopolyploids and niche differentiation can promote establishment. Despite this, the direct effects of polyploidy at the population level remain largely untested despite establishment being an intrinsically population-level process. We conducted population-level experiments where life-history investment in current and future growth was tracked in four lineage pairs of diploids and synthetic autotetraploids of the aquatic plant Spirodela polyrhiza. Population growth was evaluated with and without competition between diploids and neopolyploids across a range of nutrient treatments. Although neopolyploid populations produce more biomass, they reach lower population sizes and have reduced carrying capacities when growing alone or in competition across all nutrient treatments. Thus, contrary to individual-level studies, our population-level data suggest that neopolyploids are competitively inferior to diploids. Conversely, neopolyploid populations have greater investment in dormant propagule production than diploids. Our results show that neopolyploid populations should not persist based on current growth dynamics, but high potential future growth may allow polyploids to establish in subsequent seasons.

Plant neopolyploidy and genetic background differentiate the microbiome of duckweed across a variety of natural freshwater sources.

Whole-genome duplication has long been appreciated for its role in driving phenotypic novelty in plants, often altering the way organisms interface with the abiotic environment. Only recently, however, have we begun to investigate how polyploidy influences interactions of plants with other species, despite the biotic niche being predicted as one of the main determinants of polyploid establishment. Nevertheless, we lack information about how polyploidy affects the diversity and composition of the microbial taxa that colonize plants, and whether this is genotype-dependent and repeatable across natural environments. This information is a first step towards understanding whether the microbiome contributes to polyploid establishment. We, thus, tested the immediate effect of polyploidy on the diversity and composition of the bacterial microbiome of the aquatic plant Spirodela polyrhiza using four pairs of diploids and synthetic autotetraploids. Under controlled conditions, axenic plants were inoculated with pond waters collected from 10 field sites across a broad environmental gradient. Autotetraploids hosted 4%-11% greater bacterial taxonomic and phylogenetic diversity than their diploid progenitors. Polyploidy, along with its interactions with the inoculum source and genetic lineage, collectively explained 7% of the total variation in microbiome composition. Furthermore, polyploidy broadened the core microbiome, with autotetraploids having 15 unique bacterial taxa in addition to the 55 they shared with diploids. Our results show that whole-genome duplication directly leads to novelty in the plant microbiome and importantly that the effect is dependent on the genetic ancestry of the polyploid and generalizable over many environmental contexts.

The pollen virome: A review of pollen-associated viruses and consequences for plants and their interactions with pollinators.

The movement of pollen grains from anthers to stigmas, often by insect pollinator vectors, is essential for plant reproduction. However, pollen is also a unique vehicle for viral spread. Pollen-associated plant viruses reside on the outside or inside of pollen grains, infect susceptible individuals through vertical or horizontal infection pathways, and can decrease plant fitness. These viruses are transferred with pollen between plants by pollinator vectors as they forage for floral resources; thus, pollen-associated viral spread is mediated by floral and pollen grain phenotypes and pollinator traits, much like pollination. Most of what is currently known about pollen-associated viruses was discovered through infection and transmission experiments in controlled settings, usually involving one virus and one plant species of agricultural or horticultural interest. In this review, we first provide an updated, comprehensive list of the recognized pollen-associated viruses. Then, we summarize virus, plant, pollinator vector, and landscape traits that can affect pollen-associated virus transmission, infection, and distribution. Next, we highlight the consequences of plant-pollinator-virus interactions that emerge in complex communities of co-flowering plants and pollinator vectors, such as pollen-associated virus spread between plant species and viral jumps from plant to pollinator hosts. We conclude by emphasizing the need for collaborative research that bridges pollen biology, virology, and pollination biology.

Interspecific variation in resistance and tolerance to herbicide drift reveals potential consequences for plant community co-flowering interactions and structure at the agro-eco interface.

BACKGROUND AND AIMS: When plant communities are exposed to herbicide 'drift', wherein particles containing the active ingredient travel off-target, interspecific variation in resistance or tolerance may scale up to affect community dynamics. In turn, these alterations could threaten the diversity and stability of agro-ecosystems. We investigated the effects of herbicide drift on the growth and reproduction of 25 wild plant species to make predictions about the consequences of drift exposure on plant-plant interactions and the broader ecological community. METHODS: We exposed potted plants from species that commonly occur in agricultural areas to a drift-level dose of the widely used herbicide dicamba or a control solution in the glasshouse. We evaluated species-level variation in resistance and tolerance for vegetative and floral traits. We assessed community-level impacts of drift by comparing the species evenness and flowering networks of glasshouse synthetic communities comprised of drift-exposed and control plants. KEY RESULTS: Species varied significantly in resistance and tolerance to dicamba drift: some were negatively impacted while others showed overcompensatory responses. Species also differed in the way they deployed flowers over time following drift exposure. While drift had negligible effects on community evenness based on vegetative biomass, it caused salient differences in the structure of co-flowering networks within communities. Drift reduced the degree and intensity of flowering overlap among species, altered the composition of groups of species that were more likely to co-flower with each other than with others and shifted species roles (e.g. from dominant to inferior floral producers, and vice versa). CONCLUSIONS: These results demonstrate that even low levels of herbicide exposure can significantly alter plant growth and reproduction, particularly flowering phenology. If field-grown plants respond similarly, then these changes would probably impact plant-plant competitive dynamics and potentially plant-pollinator interactions occurring within plant communities at the agro-ecological interface.

Herbicides as anthropogenic drivers of eco-evo feedbacks in plant communities at the agro-ecological interface.

Herbicides act as human-mediated novel selective agents and community disruptors, yet their full effects on eco-evolutionary dynamics in natural communities have only begun to be appreciated. Here, we synthesize how herbicide exposures can result in dramatic phenotypic and compositional shifts within communities at the agro-ecological interface and how these in turn affect species interactions and drive plant (and plant-associates') evolution in ways that can feedback to continue to affect the ecology and ecosystem functions of these assemblages. We advocate a holistic approach to understanding these dynamics that includes plastic changes and plant community transformations and also extends beyond this single trophic level targeted by herbicides to the effects on nontarget plant-associated organisms and their potential to evolve, thereby embracing the complexity of these real-world systems. We make explicit recommendations for future research to achieve this goal and specifically address impacts of ecology on evolution, evolution on ecology and their feedbacks so that we can gain a more predictive view of the fates of herbicide-impacted communities.

Pollinators mediate floral microbial diversity and microbial network under agrochemical disturbance.

How pollinators mediate microbiome assembly in the anthosphere is a major unresolved question of theoretical and applied importance in the face of anthropogenic disturbance. We addressed this question by linking visitation of diverse pollinator functional groups (bees, wasps, flies, butterflies, beetles, true bugs and other taxa) to the key properties of the floral microbiome (microbial α- and ß-diversity and microbial network) under agrochemical disturbance, using a field experiment of bactericide and fungicide treatments on cultivated strawberries that differ in flower abundance. Structural equation modelling was used to link agrochemical disturbance and flower abundance to pollinator visitation to floral microbiome properties. Our results revealed that (i) pollinator visitation influenced the α- and ß-diversity and network centrality of the floral microbiome, with different pollinator functional groups affecting different microbiome properties; (ii) flower abundance influenced the floral microbiome both directly by governing the source pool of microbes and indirectly by enhancing pollinator visitation; and (iii) agrochemical disturbance affected the floral microbiome primarily directly by fungicide, and less so indirectly via pollinator visitation. These findings improve the mechanistic understanding of floral microbiome assembly, and may be generalizable to many other plants that are visited by diverse insect pollinators in natural and managed ecosystems.

Repeated translocation of a gene cassette drives sex-chromosome turnover in strawberries.

Turnovers of sex-determining systems represent important diversifying forces across eukaryotes. Shifts in sex chromosomes-but conservation of the master sex-determining genes-characterize distantly related animal lineages. Yet in plants, in which separate sexes have evolved repeatedly and sex chromosomes are typically homomorphic, we do not know whether such translocations drive sex-chromosome turnovers within closely related taxonomic groups. This phenomenon can only be demonstrated by identifying sex-associated nucleotide sequences, still largely unknown in plants. The wild North American octoploid strawberries (Fragaria) exhibit separate sexes (dioecy) with homomorphic, female heterogametic (ZW) inheritance, yet sex maps to three different chromosomes in different taxa. To characterize these turnovers, we identified sequences unique to females and assembled their reads into contigs. For most octoploid Fragaria taxa, a short (13 kb) sequence was observed in all females and never in males, implicating it as the sex-determining region (SDR). This female-specific "SDR cassette" contains both a gene with a known role in fruit and pollen production and a novel retrogene absent on Z and autosomal chromosomes. Phylogenetic comparison of SDR cassettes revealed three clades and a history of repeated translocation. Remarkably, the translocations can be ordered temporally due to the capture of adjacent sequence with each successive move. The accumulation of the "souvenir" sequence-and the resultant expansion of the hemizygous SDR over time-could have been adaptive by locking genes into linkage with sex. Terminal inverted repeats at the insertion borders suggest a means of movement. To our knowledge, this is the first plant SDR shown to be translocated, and it suggests a new mechanism ("move-lock-grow") for expansion and diversification of incipient sex chromosomes.

Pollinator effectiveness is affected by intraindividual behavioral variation.

Variation in pollinator quality is fundamental to the evolution of plant-pollinator mutualisms and such variation frequently results from differences in foraging behavior. Surprisingly, despite substantial intraindividual variation in pollinator foraging behavior, the consequences for pollen removal and deposition on flowers are largely unknown. We asked how two pollen foraging behaviors of a generalist pollinator (Bombus impatiens) affect removal and deposition of heterospecific and conspecific pollen, key aspects of pollinator quality, for multiple plant species. In addition, we examined how bee body size and pollen placement among body parts shaped pollen movement. We manipulated foraging behavior types using artificial flowers, which donated pollen that captive bees then deposited on three recipient plant species. While body size primarily affected donor pollen removal, foraging behavior primarily affected donor pollen deposition. How behavior affected donor pollen deposition depended on the plant species and the quantity of donor pollen on the bee's abdomen. Plant species with smaller stigmas received significantly less pollen and fewer bees successfully transferred pollen to them. For a single plant species, heterospecific pollen interfered with conspecific pollen deposition, such that more heterospecific pollen on the bee's abdomen resulted in less conspecific pollen deposition on the flower. Thus, intraindividual variation in foraging behavior and its interaction with the amount and placement of acquired pollen and with floral morphology can affect pollinator quality and may shape plant fitness via both conspecific and heterospecific pollen transfer.

Diversity and composition of pollen loads carried by pollinators are primarily driven by insect traits, not floral community characteristics.

Flowering plants require conspecific pollen to reproduce but they often also receive heterospecific pollen, suggesting that pollinators carry mixed pollen loads. However, little is known about drivers of abundance, diversity or composition of pollen carried by pollinators. Are insect-carried pollen loads shaped by pollinator traits, or do they reflect available floral resources? We quantified pollen on 251 individual bees and 95 flies in a florally diverse community. We scored taxonomic order, sex, body size, hairiness and ecological specialization of pollinators, and recorded composition of available flowers. We used phylogenetically controlled model selection to compare relative influences of pollinator traits and floral resources on abundance, diversity and composition of insect-carried pollen. We tested congruence between composition of pollen loads and available flowers. Pollinator size, specialization and type (female bee, male bee, or fly) described pollen abundance, diversity and composition better than floral diversity. Pollen loads varied widely among insects (10-80,000,000 grains, 1-16 species). Pollen loads of male bees were smaller, but vastly more diverse than those of female bees, and equivalent in size but modestly more diverse than those of flies. Pollen load size and diversity were positively correlated with body size but negatively correlated with insect ecological specialization. These traits also drove variation in taxonomic and phylogenetic composition of insect-carried pollen loads, but composition was only weakly congruent with available floral resources. Qualities of pollinators best predict abundance and diversity of carried pollen indicating that functional composition of pollinator communities may be important to structuring heterospecific pollen transfer among plants.

Polyploid plants obtain greater fitness benefits from a nutrient acquisition mutualism.

Polyploidy is a key driver of ecological and evolutionary processes in plants, yet little is known about its effects on biotic interactions. This gap in knowledge is especially profound for nutrient acquisition mutualisms, despite the fact that they regulate global nutrient cycles and structure ecosystems. Generalism in mutualistic interactions depends on the range of potential partners (niche breadth), the benefits obtained and ability to maintain benefits across a variety of partners (fitness plasticity). Here, we determine how each of these is influenced by polyploidy in the legume-rhizobium mutualism. We inoculated a broad geographic sample of natural diploid and autotetraploid alfalfa (Medicago sativa) lineages with a diverse panel of Sinorhizobium bacterial symbionts. To analyze the extent and mechanism of generalism, we measured host growth benefits and functional traits. Autotetraploid plants obtained greater fitness enhancement from mutualistic interactions and were better able to maintain this across diverse rhizobial partners (i.e. low plasticity in fitness) relative to diploids. These benefits were not attributed to increases in niche breadth, but instead reflect increased rewards from investment in the mutualism. Polyploid plants displayed greater generalization in bacterial mutualisms relative to diploids, illustrating another axis of advantage for polyploids over diploids.

TRY plant trait database - enhanced coverage and open access.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

Pollen on stigmas as proxies of pollinator competition and facilitation: complexities, caveats and future directions.

BACKGROUND: Pollen transfer via animals is necessary for reproduction by ~80 % of flowering plants, and most of these plants live in multispecies communities where they can share pollinators. While diffuse plant-pollinator interactions are increasingly recognized as the rule rather than the exception, their fitness consequences cannot be deduced from flower visitation alone, so other proxies, functionally closer to seed production and amenable for use in a broad variety of diverse communities, are necessary. SCOPE: We conceptually summarize how the study of pollen on stigmas of spent flowers can reflect key drivers and functional aspects of the plant-pollinator interaction (e.g. competition, facilitation or commensalism). We critically evaluate how variable visitation rates and other factors (pollinator pool and floral avoidance) can give rise to different relationships between heterospecific pollen and (1) conspecific pollen on the stigma and (2) conspecific tubes/grain in the style, revealing the complexity of potential interpretations. We advise on best practices for using these proxies, noting the assumptions and caveats involved in their use, and explicate what additional data are required to verify interpretation of given patterns. CONCLUSIONS: We conclude that characterizing pollen on stigmas of spent flowers provides an attainable indirect measure of pollination interactions, but given the complex processes of pollen transfer that generate patterns of conspecific-heterospecific pollen on stigmas these cannot alone determine whether competition or facilitation are the underlying drivers. Thus, functional tests are also needed to validate these hypotheses.

Autopolyploidy alters nodule-level interactions in the legume-rhizobium mutualism.

PREMISE: Polyploidy is a major genetic driver of ecological and evolutionary processes in plants, yet its effects on plant interactions with mutualistic microbes remain unresolved. The legume-rhizobium symbiosis regulates global nutrient cycles and plays a role in the diversification of legume species. In this mutualism, rhizobia bacteria fix nitrogen in exchange for carbon provided by legume hosts. This exchange occurs inside root nodules, which house bacterial cells and represent the interface of legume-rhizobium interactions. Although polyploidy may directly impact the legume-rhizobium mutualism, no studies have explored how it alters the internal structure of nodules. METHODS: We created synthetic autotetraploids using Medicago sativa subsp. caerulea. Neotetraploid plants and their diploid progenitors were singly inoculated with two strains of rhizobia, Sinorhizobium meliloti and S. medicae. Confocal microscopy was used to quantify internal traits of nodules produced by diploid and neotetraploid plants. RESULTS: Autotetraploid plants produced larger nodules with larger nitrogen fixation zones than diploids for both strains of rhizobia, although the significance of these differences was limited by power. Neotetraploid M. sativa subsp. caerulea plants also produced symbiosomes that were significantly larger, nearly twice the size, than those present in diploids. CONCLUSIONS: This study sheds light on how polyploidy directly affects a plant-bacterium mutualism and uncovers novel mechanisms. Changes in plant-microbe interactions that directly result from polyploidy likely contribute to the increased ability of polyploid legumes to establish in diverse environments.

Genome duplication effects on functional traits and fitness are genetic context and species dependent: studies of synthetic polyploid Fragaria.

PREMISE: Divergence in functional traits and adaptive responses to environmental change underlies the ecological advantage of polyploid plants in the wild. While established polyploids may benefit from combined outcomes of genome doubling, hybridization, and polyploidy-enabled adaptive evolution, whether genome doubling alone can drive ecological divergence or whether the outcome is genetically variable remains less clear. METHODS: Using synthetic, colchicine-induced, autotetraploid (4x) plants derived from self-pollinated diploid (2x) seeds, and their colchicine-treated but unconverted diploid (2x.nc) full sibs from two diploid wild strawberry taxa (Fragaria vesca subsp. vesca and F. vesca subsp. bracteata), we examined the effects of genome doubling on functional traits, heat stress tolerance, and fitness components across taxa and maternal families (i.e., genetic families) within taxa. RESULTS: Comparisons between 2x and 2x.nc plants indicated a negligible effect of colchicine treatment on functional traits. Genome doubling increased stomatal length and decreased stomatal density, specific leaf area, and leaf vein density, recapitulating patterns observed in wild polyploid Fragaria. Trichome density, heat stress tolerance, and relative growth rate were not significantly affected by genome doubling. Although clonal reproduction was reduced in response to genome doubling, this effect was strongly genetic-family dependent. CONCLUSIONS: The results suggest that genome doubling during incipient speciation alone can generate ecological divergence and variation among genetic lineages. This response potentially allows for rapid short-term evolutionary adaptation and fuels genomic diversity and independent origins of polyploidy.

Pollen on Stigmas of Herbarium Specimens: A Window into the Impacts of a Century of Environmental Disturbance on Pollen Transfer.

Pollination is necessary for plant reproduction but often highly susceptible to disruption, for example, by habitat fragmentation and climate change. Here, we indirectly evaluated on a century timescale pollination interactions for species in one of the historically most disturbed habitats on earth-tropical dry forests of Hawai'i. We employed a novel method for acquiring a historical perspective on temporal change in pollination by characterizing pollen on stigmas of herbarium specimens from six remnant native species collected from 1909-2002. We determine whether temporal shifts occurred in (1) pollination quantity and quality or (2) the composition of species interacting via pollen transfer. While pollen quantity remained constant, these remnant species interact with different species in modern times via pollen transfer than they did nearly 100 years ago. Species that are resilient to long-term environmental change may also be the ones subject to changes in pollination interactions.

Consequences of invasion for pollen transfer and pollination revealed in a tropical island ecosystem.

Pollination is known to be sensitive to environmental change but we lack direct estimates of how quantity and quality of pollen transferred between plant species shifts along disturbance gradients. This limits our understanding of how species compositional change impacts pollen receipt per species and structure of pollen transfer networks. We constructed pollen transfer networks along a plant invasion gradient in the Hawaiian dry tropical forest ecosystem. Flowers and stigmas were collected from both native and introduced plants, pollen was identified and enumerated and floral traits were measured. We also characterized pollen loads carried by individuals of the dominant invasive pollinator, Apis mellifera. Species flowering in native-dominated sites were more tightly connected by pollen transfer than those in heavily invaded sites. Compositional turnover in the pollen loads of A. mellifera was correlated (70%) with turnover in the composition of pollen transfer networks. Floral traits predicted species roles within pollen transfer networks, but many of these differed qualitatively depending on whether plants were native or introduced. Our work indicates that pollen transfer networks change with invasion. Floral morphology and foraging behaviour of the introduced super-generalist pollinator are implicated as key in determining the roles introduced species play within native pollen transfer networks.