Academic literature on the topic 'Generalist pollinators'

AbstractWe consider the role of generalist Diptera in the pollination of two dioecious plant species, Clematis ligusticifolia Nutt. (Ranunculaceae) and Shepherdia canadensis (L.) Nutt. (Elaeagnaceae). In particular, we assess (i) whether or not generalist pollinators are unable to distinguish between the sexes of dioecious species and so visit the sexes at equivalent rates, and (ii) the number of flowers that generalist flies visit and revisit during a foraging bout, which would affect self-pollination if plants were hermaphroditic. We determined the visitation rate to each plant species during 10 min periods and recorded the number of flowers that individual pollinators visited and revisited per foraging bout. Diptera were the main pollinators, visiting both sexes at similar rates for both plant species. The main visitors to C. ligusticifolia were muscoid flies (small and large), Culicidae, and halictid bees. The number of flowers visited in this species varied with pollinator group, but groups did not differ in the frequency of revisits. Visitors to S. canadensis were primarily Syrphidae and Empididae. Neither the number of flowers visited nor the number of revisits differed between these two pollinator groups. The results for each plant species are discussed and contrasted, particularly with other studies of the behaviour of generalist and specialist pollinators. We compare the observed pollinator behaviours, and their implications for plant mating, with the various theories of the role of pollinators in the evolution of the dioecious breeding system in plants.

A generalist pollination system may be characterized through the interaction of a plant species with two or more functional groups of pollinators. The spatiotemporal variation of the most effective pollinator is the factor most frequently advocated to explain the emergence and maintenance of generalist pollination systems. There are few studies merging variation in floral visitor assemblages and the efficacy of pollination by different functional groups. Thus, there are gaps in our knowledge about the variation in time of pollinator efficacy and frequency of generalist species. In this study, we evaluated the pollination efficacy of the floral visitors of Edmundoa lindenii (Bromeliaceae) and their frequency of visits across four reproductive events. We analyzed the frequency of the three groups of floral visitors (large bees, small bees, and hummingbirds) through focal observations in the reproductive events of 2015, 2016, 2017, and 2018. We evaluated the pollination efficacy (fecundity after one visit) through selective exposure treatments and the breeding system by manual pollinations. We tested if the reproductive success after natural pollination varied between the reproductive events and also calculated the pollen limitation index. E. lindenii is a self-incompatible and parthenocarpic species, requiring the action of pollinators for sexual reproduction. Hummingbirds had higher efficacy than large bees and small bees acted only as pollen larcenists. The relative frequency of the groups of floral visitors varied between the reproductive events. Pollen limitation has occurred only in the reproductive event of 2017, when visits by hummingbirds were scarce and reproductive success after natural pollination was the lowest. We conclude that hummingbirds and large bees were the main and the secondary pollinators of E. lindenii, respectively, and that temporal variations in the pollinator assemblages had effects on its reproductive success. Despite their lower pollination efficacy, large bees ensured seed set when hummingbirds failed. Thus, we provide evidence that variable pollination environments may favor generalization, even under differential effectiveness of pollinator groups if secondary pollinators provide reproductive assurance.

The structural patterns comprising bimodal pollination networks can help characterize plant–pollinator systems and the interactions that influence species distribution and diversity over time and space. We compare network organization of three plant–pollinator communities along the altitudinal gradient of the San Francisco Peaks in northern Arizona. We found that pollination networks become more nested, as well as exhibit lower overall network specialization, with increasing elevation. Greater weight of generalist pollinators at higher elevations of the San Francisco Peaks may result in plant–pollinator communities less vulnerable to future species loss due to changing climate or shifts in species distribution. We uncover the critical, more generalized pollinator species likely responsible for higher nestedness and stability at the higher elevation environment. The generalist species most important for network stability may be of the greatest interest for conservation efforts; preservation of the most important links in plant–pollinator networks may help secure the more specialized pollinators and maintain species redundancy in the face of ecological change, such as changing climate.

Interactions between pollinators and their plant hosts are central to maintaining global biodiversity and ensuring our food security. In this special issue, we compile reviews that summarize existing knowledge and point out key outstanding research areas to understand and safeguard pollinators, pollinators–host plant interactions and the pollination ecosystem services they provide. The vast diversity of the pollinator–plant interactions that exists on this planet still remains poorly explored, with many being associations involving a specialist pollinator partner, although historically most focus has been given to generalist pollinators, such as the honeybee. Two areas highlighted here are the ecology and evolution of oligolectic bee species, and the often-neglected groups of pollinators that forage solely at night. Advances in automated detection technologies could offer potential and complementary solutions to the current shortfall in knowledge on interactions occurring between less well-documented plant–pollinator associations, by increasing the collection range and capacity of flower visitation data over space and time. Pollinator–host plant interactions can be affected by external biotic factors, with herbivores and pathogens playing particularly important roles. Such interactions can be disrupted by modifying plant volatile and reward chemistry, with possible effects on pollinator attraction and pollination success. Mechanisms which underpin interactions between plants and their pollinators also face many anthropogenic disturbances. Reviews in this issue discuss threats from parasites and climate change to pollinator populations and plant–pollinator networks, and suggest new ways to mitigate these threats. While the protection of existing plant–pollinator networks will be a crucial goal for conservation biology, more research is needed to understand how lost interactions in degraded habitats may be restored with mutual benefits to plants and pollinators.

With more than 80% of flowering plant species specialized for animal pollination, understanding how wild pollinators utilize resources across environments can encourage efficient planting and maintenance strategies to maximize pollination and establish resilience in the face of environmental change. A fundamental question is how generalist pollinators recognize “flower objects” in vastly different ecologies and environments. On one hand, pollinators could employ a specific set of floral cues regardless of environment. Alternatively, wild pollinators could recognize an exclusive signature of cues unique to each environment or flower species. Hoverflies, which are found across the globe, are one of the most ecologically important alternative pollinators after bees and bumblebees. Here, we have exploited their cosmopolitan status to understand how wild pollinator preferences change across different continents. Without employing any a priori assumptions concerning the floral cues, we measured, predicted, and finally artificially recreated multimodal cues from individual flowers visited by hoverflies in three different environments (hemiboreal, alpine, and tropical) using a field-based methodology. We found that although “flower signatures” were unique for each environment, some multimodal lures were ubiquitously attractive, despite not carrying any reward, or resembling real flowers. While it was unexpected that cue combinations found in real flowers were not necessary, the robustness of our lures across insect species and ecologies could reflect a general strategy of resource identification for generalist pollinators. Our results provide insights into how cosmopolitan pollinators such as hoverflies identify flowers and offer specific ecologically based cues and strategies for attracting pollinators across diverse environments.

Flowering plants that attract a diverse range of pollinators represent a generalist pollination system. Studying these plants provides valuable information about accessibility of floral resources to pollinators, which is particularly important in areas where scarcity of flowers limits pollinator populations. Here, we describe the flowering phenology, reproductive biology, and visitor community of Cleomella serrulata (Pursh) Roalson & J.C.Hall and Polanisia dodecandra (L.) DC., two native species with generalist pollination systems and limited distribution in Albertan prairies. Although their flowers are similar, they differ in traits such as petal colour, inflorescence size, and nectar display. Both species were facultatively cross-pollinated and exhibited nocturnal anthesis but differed in nectar production patterns. Cleomella serrulata produced highest nectar volume in the morning and highest sugar concentration at noon, while Polanisia dodecandra produced highest nectar volume before noon but sugar concentrations were higher at sunset. We observed 150 insect taxa visiting the plants, with Hymenoptera and Diptera as the most frequent visitors for Cleomella serrulata and Polanisia dodecandra, respectively. We recorded the first nocturnal flower visitors for Cleomella serrulata and the first record of Nysson plagiatus (Cresson) for Alberta. Both plant species present effective nectar and pollen resources for pollinators at the study sites and may be useful in the maintenance of native pollinators in at-risk prairie ecosystems.

Abstract:The ability of plants to colonize new habitats is influenced by their dependence on effective pollinators. This can be very important for plants that require specialized pollinators, especially when they disperse to islands that have low pollinator diversity. One form of specialization involves plants that require buzz-pollination, where bees must vibrate poricidal anthers at frequencies that allow pollen to be released. Pollen larceny is a phenomenon where insects ‘steal’ pollen from flowers which usually results in reduced pollination, but in some cases there can be a small contribution to pollination. Here we report pollen larceny in an endemic Fijian halictine bee Homalictus fijiensis that steals pollen by chewing anthers of the invasive weed Solanum torvum, which is a pollen-only plant requiring buzz pollination. In over nine hours of observations at six sites where H. fijiensis visited S. torvum, it never attempted to locate nectaries, it never buzzed anthers, and instead chewed anther tips, indicating an adaptation to exploit nectarless flowers with poricidal anthers without buzz-pollination. Analyses of 30 pollen loads from H. fijiensis collected from S. torvum flowers indicate 27 of these contained S. torvum pollen, ranging from 1% to 99% of total pollen, indicating it is a pollen vector for this plant. Our findings support arguments that super-generalist pollinators in island ecosystems can promote the spread of invasive plants, but go further by indicating that super-generalist strategies can extend to plants with highly specialized pollinator requirements.

The reciprocal importance of bromeliads and hummingbirds has been proposed for many years, even suggesting coevolution between these two groups. Nevertheless, data are lacking that allow a better test of the relationships involved. Here we investigate the relationship between bromeliads and hummingbirds in an area of secondary Atlantic rain forest in southern Brazil. The study examined the interactions among 12 species of bromeliad and 10 of hummingbird at Reserva Natural Salto Morato, Paraná state. The number of flowering species of bromeliad and the species richness and abundance of hummingbirds were quantified monthly between November 2004 and October 2005. Focal observations on each bromeliad species were made to determine the hummingbird visitors. Neither species richness nor abundance of hummingbirds were related to bromeliad phenology. Together with the monthly variation in visit frequency by a given pollinator to a given plant, these factors indicate a generalization in the use of bromeliads by hummingbirds and argue against tight coevolution. Ramphodon naevius and Thalurania glaucopis were the main pollinators in the community. Aechmea nudicaulis was the most generalist bromeliad species. The generalist species interacted with other generalists or with asymmetric specialists and there was no specialist–specialist interaction. This produced a strongly organized and nested matrix of interactions. This nestedness is similar to other plant-pollinators networks, supporting the hypothesis that the evolutionary relationship between bromeliads and hummingbirds is no stronger than that of other pollination networks.

Abstract In the context of global pollinator decline, little is known about the protection status and ecology of many species. This lack of knowledge is particularly important for Mediterranean protected areas that harbor diverse pollinator communities and are subject to considerable anthropogenic pressures. Calanques National Park (85 km2), which is located near Marseille (France), is dominated by Mediterranean low-vegetation habitats, such as phrygana and scrublands. These habitats offer favorable conditions for pollinator species due to the important amount of floral resources. Within a 10-yr period, we recorded bee (Hymenoptera: Apoidea: Anthophila), hover fly (Diptera: Syrphidae), and bee fly (Diptera: Bombyliidae) species and their interactions with the local flora through 10 field campaigns. We caught 250 pollinator species, including 192 bees, 38 hover flies, and 20 bee flies, for a total of 2,770 specimens. We recorded seven threatened bees (six near threatened and one endangered). Among the bee species, 47.9% were below-ground nesting species, and 54.7% were generalist species. Analysis of the pollination network showed that generalist and specialist pollinators do not share the same floral resources. The Cistaceae plant family (Malvales: Cistaceae) acted as a central node in the plant–pollinator network, interacting with 52 different pollinator species, which shows the importance of large open flowers that could be easily visited by both short and long-tongued pollinators in Mediterranean habitats. The occurrence of pollinator species and their ecological traits should strongly contribute to reinforcing the available information to provide or ameliorate the conservation statuses determined by IUCN Red List.

Abstract Communication is often vital to the maintenance of mutualisms. In plant-pollinator mutualisms, plants signal pollinators via floral displays, composed of olfactory, visual, and other plant-derived cues. While plants are understood to be associated with microbes, only recently has the role of microbial (yeast and bacteria) inhabitants of flowers as intermediaries of plant-pollinator communication been recognized. Animals frequently use microbial cues to find resources, yet no study has examined whether microbes directly mediate learned and innate pollinator responses. Here, we asked whether microbes on the flower surface, independent of their modification of floral rewards, can mediate these key components of pollinator preference. In the field, we characterized flower and bumble bee microbial abundance, and in laboratory assays we tested whether bumble bees (Bombus impatiens) discriminated flowers on the basis of an experimental floral microbial community on the petals and whether microbe-derived chemicals were effective cues. Learning of microbial community cues was associative and reward context-dependent and mediated by microbial chemicals. Deconstructing the experimental microbial community showed bees innately avoided flowers with bacteria, but were undeterred by yeast. Microbial cues thus potentially facilitate dynamic communication between plants and pollinators such as bumble bees, especially as pollinator visitation can change flower microbiota. We suggest that the study of communication in mutualism generally would benefit by considering not only the multicellular eukaryote partners, but their microbial associates.

A renewed focus on generalised pollinator systems has inspired a conceptual framework which highlights that spatial and temporal interactions among plants and their assemblage of pollinators can vary across the individual, population, regional and species levels. Pollination is clearly a dynamic interaction, varying in the number and interdependence of participants and the strength of the outcome of the interaction. Therefore, the role of variation in pollination is fundamental for understanding ecological dynamics of plant populations and is a major factor in the evolution and maintenance of generalised and specialised pollination systems. My study centred on these basic concepts by addressing the following questions: (1) How variable are pollinators in a generalised pollination system? To what degree do insect visitation rates and assemblage composition vary spatially among populations and temporally among flowering seasons? (2) How does variation in pollinators affect plant reproductive success? I chose to do this using a model system, Trachymene incisa subsp. incisa (Apiaceae), which is a widespread Australian herbaceous species with simple white flowers grouped into umbels that attract a high diversity of insect visitors. The Apiaceae are considered to be highly generalist in terms of pollination, due to their simple and uniform floral display and easily accessible floral rewards. Three populations of T. incisa located between 70 km and 210 km apart were studied over 2-3 years. The few studies investigating spatial and temporal variation simultaneously over geographic and yearly/seasonal scales indicate that there is a trend for more spatial than temporal variation in pollinators of generalist-pollinated plants. My study showed both spatial and temporal variation in assemblage composition among all populations and variation in insect visitation rates, in the form of a significant population by year interaction. However, removing ants from the analyses to restrict the assemblage to flying insects and the most likely pollinators, resulted in a significant difference in overall visitation rate between years but no difference in assemblage composition between the Myall Lakes and Tomago populations. These results indicate more temporal than spatial variation in the flying insect visitor assemblage of T. incisa. Foraging behaviour provides another source of variation in plant-pollinator interactions. Trachymene incisa exhibits umbels that function as either male or female at any one time and offer different floral rewards in each phase. For successful pollination, pollinators must visit both male and female umbels during a foraging trip. Insects showed both preferences and non-preferences for umbel phases in natural patches where the gender ratio was male biased. In contrast, insects showed no bias in visitation during a foraging trip or in time spent foraging on male and female umbels in experimental arrays where the gender ratio was equal. Pollinator assemblages consisting of a mixture of different pollinator types coupled with temporal variation in the assemblages of populations among years maintains generalisation at the population/local level. In addition, spatial variation in assemblages among populations maintains generalisation at the species level. Fire alters pollination in T. incisa by shifting the flowering season and reducing the abundance of flying insects. Therefore, fire plays an important role in maintaining spatial and temporal variation in this fire-prone system. Although insect pollinators are important in determining the mating opportunities of 90% of flowering plant species worldwide, few studies have looked at the effects of variation in pollinator assemblages on plant reproductive success and mating. In T. incisa, high insect visitation rates do not guarantee high plant reproductive success, indicating that the quality of visit is more important than the rate of visitation. This is shown by comparing the Agnes Banks and Myall Lakes populations in 2003: Agnes Banks received the highest visitation rate from an assemblage dominated by ants but produced the lowest reproductive output, and Myall Lakes received the lowest visitation rate by an assemblage dominated by a native bee and produced the highest seedling emergence. Interestingly, populations with different assemblage composition can produce similar percentage seed set per umbel. However, similar percentage seed set did not result in similar percentage seedling emergence. Differences among years in reproductive output (total seed production) were due to differences in umbel production (reproductive effort) and proportion of umbels with seeds, and not seed set per umbel. Trachymene incisa is self-compatible and suffers weak to intermediate levels of inbreeding depression through early stages of the life cycle when seeds are self-pollinated and biparentally inbred. Floral phenology, in the form of synchronous protandry, plays an important role in avoiding self-pollination within umbels and reducing the chance of geitonogamous pollination between umbels on the same plant. Although pollinators can increase the rate of inbreeding in T. incisa by foraging on both male and female phase umbels on the same plant or closely related plants, most consecutive insect movements were between plants not located adjacent to each other. This indicates that inbreeding is mostly avoided and that T. incisa is a predominantly outcrossing species, although further genetic analyses are required to confirm this hypothesis. A new conceptual understanding has emerged from the key empirical results in the study of this model generalised pollination system. The large differences among populations and between years indicate that populations are not equally serviced by pollinators and are not equally generalist. Insect visitation rates varied significantly throughout the day, highlighting that sampling of pollinators at one time will result in an inaccurate estimate and usually underestimate the degree of generalisation. The visitor assemblage is not equivalent to the pollinator assemblage, although non-pollinating floral visitors are likely to influence the overall effectiveness of the pollinator assemblage. Given the high degree of variation in both the number of pollinator species and number of pollinator types, I have constructed a model which includes the degree of ecological and functional specialisation of a plant species on pollinators and the variation encountered across different levels of plant organisation. This model describes the ecological or current state of plant species and their pollinators, as well as presenting the patterns of generalisation across a range of populations, which is critical for understanding the evolution and maintenance of the system. In-depth examination of pollination systems is required in order to understand the range of strategies utilised by plants and their pollinators, and I advocate a complete floral visitor assemblage approach to future studies in pollination ecology. In particular, future studies should focus on the role of introduced pollinators in altering generalised plant-pollinator systems and the contribution of non-pollinating floral visitors to pollinator assemblage effectiveness. Comparative studies involving plants with highly conserved floral displays, such as those in the genus Trachymene and in the Apiaceae, will be useful for investigating the dynamics of generalised pollination systems across a range of widespread and restricted species.

Doctor of Philosophy (PhD)
A renewed focus on generalised pollinator systems has inspired a conceptual framework which highlights that spatial and temporal interactions among plants and their assemblage of pollinators can vary across the individual, population, regional and species levels. Pollination is clearly a dynamic interaction, varying in the number and interdependence of participants and the strength of the outcome of the interaction. Therefore, the role of variation in pollination is fundamental for understanding ecological dynamics of plant populations and is a major factor in the evolution and maintenance of generalised and specialised pollination systems. My study centred on these basic concepts by addressing the following questions: (1) How variable are pollinators in a generalised pollination system? To what degree do insect visitation rates and assemblage composition vary spatially among populations and temporally among flowering seasons? (2) How does variation in pollinators affect plant reproductive success? I chose to do this using a model system, Trachymene incisa subsp. incisa (Apiaceae), which is a widespread Australian herbaceous species with simple white flowers grouped into umbels that attract a high diversity of insect visitors. The Apiaceae are considered to be highly generalist in terms of pollination, due to their simple and uniform floral display and easily accessible floral rewards. Three populations of T. incisa located between 70 km and 210 km apart were studied over 2-3 years. The few studies investigating spatial and temporal variation simultaneously over geographic and yearly/seasonal scales indicate that there is a trend for more spatial than temporal variation in pollinators of generalist-pollinated plants. My study showed both spatial and temporal variation in assemblage composition among all populations and variation in insect visitation rates, in the form of a significant population by year interaction. However, removing ants from the analyses to restrict the assemblage to flying insects and the most likely pollinators, resulted in a significant difference in overall visitation rate between years but no difference in assemblage composition between the Myall Lakes and Tomago populations. These results indicate more temporal than spatial variation in the flying insect visitor assemblage of T. incisa. Foraging behaviour provides another source of variation in plant-pollinator interactions. Trachymene incisa exhibits umbels that function as either male or female at any one time and offer different floral rewards in each phase. For successful pollination, pollinators must visit both male and female umbels during a foraging trip. Insects showed both preferences and non-preferences for umbel phases in natural patches where the gender ratio was male biased. In contrast, insects showed no bias in visitation during a foraging trip or in time spent foraging on male and female umbels in experimental arrays where the gender ratio was equal. Pollinator assemblages consisting of a mixture of different pollinator types coupled with temporal variation in the assemblages of populations among years maintains generalisation at the population/local level. In addition, spatial variation in assemblages among populations maintains generalisation at the species level. Fire alters pollination in T. incisa by shifting the flowering season and reducing the abundance of flying insects. Therefore, fire plays an important role in maintaining spatial and temporal variation in this fire-prone system. Although insect pollinators are important in determining the mating opportunities of 90% of flowering plant species worldwide, few studies have looked at the effects of variation in pollinator assemblages on plant reproductive success and mating. In T. incisa, high insect visitation rates do not guarantee high plant reproductive success, indicating that the quality of visit is more important than the rate of visitation. This is shown by comparing the Agnes Banks and Myall Lakes populations in 2003: Agnes Banks received the highest visitation rate from an assemblage dominated by ants but produced the lowest reproductive output, and Myall Lakes received the lowest visitation rate by an assemblage dominated by a native bee and produced the highest seedling emergence. Interestingly, populations with different assemblage composition can produce similar percentage seed set per umbel. However, similar percentage seed set did not result in similar percentage seedling emergence. Differences among years in reproductive output (total seed production) were due to differences in umbel production (reproductive effort) and proportion of umbels with seeds, and not seed set per umbel. Trachymene incisa is self-compatible and suffers weak to intermediate levels of inbreeding depression through early stages of the life cycle when seeds are self-pollinated and biparentally inbred. Floral phenology, in the form of synchronous protandry, plays an important role in avoiding self-pollination within umbels and reducing the chance of geitonogamous pollination between umbels on the same plant. Although pollinators can increase the rate of inbreeding in T. incisa by foraging on both male and female phase umbels on the same plant or closely related plants, most consecutive insect movements were between plants not located adjacent to each other. This indicates that inbreeding is mostly avoided and that T. incisa is a predominantly outcrossing species, although further genetic analyses are required to confirm this hypothesis. A new conceptual understanding has emerged from the key empirical results in the study of this model generalised pollination system. The large differences among populations and between years indicate that populations are not equally serviced by pollinators and are not equally generalist. Insect visitation rates varied significantly throughout the day, highlighting that sampling of pollinators at one time will result in an inaccurate estimate and usually underestimate the degree of generalisation. The visitor assemblage is not equivalent to the pollinator assemblage, although non-pollinating floral visitors are likely to influence the overall effectiveness of the pollinator assemblage. Given the high degree of variation in both the number of pollinator species and number of pollinator types, I have constructed a model which includes the degree of ecological and functional specialisation of a plant species on pollinators and the variation encountered across different levels of plant organisation. This model describes the ecological or current state of plant species and their pollinators, as well as presenting the patterns of generalisation across a range of populations, which is critical for understanding the evolution and maintenance of the system. In-depth examination of pollination systems is required in order to understand the range of strategies utilised by plants and their pollinators, and I advocate a complete floral visitor assemblage approach to future studies in pollination ecology. In particular, future studies should focus on the role of introduced pollinators in altering generalised plant-pollinator systems and the contribution of non-pollinating floral visitors to pollinator assemblage effectiveness. Comparative studies involving plants with highly conserved floral displays, such as those in the genus Trachymene and in the Apiaceae, will be useful for investigating the dynamics of generalised pollination systems across a range of widespread and restricted species.

Thesis (Ph. D.)--School of Biological Sciences, Faculty of Science, University of Sydney, 2006.
Title from title screen (viewed 15 January 2009). Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy to the School of Biological Sciences, Faculty of Science. Includes bibliographical references. Also available in print form.

Plant-pollinator networks have shown to be highly dynamic systems as species and interactions change in time and space. Few studies have incorporated several habitats in their network. In this work I investigate interacting plant and pollinator communities of three adjacent habitats at Gotska Sandön, an island in the Baltic Sea. The networks varied in size between the habitats, and the larger networks of the dune and meadow displayed both nested and modular structure while the smaller forest network was more randomly organised. We found species present in more than one habitat that connected the networks by forming inter-habitat modules of tightly linked species. Species took on different topological roles in the networks depending on how many links they formed and where these attached. The habitat generalists were important to overall network structure as role correlated with habitat generalisation level.

Dissertação de mestrado em Ecologia, apresentada ao Departamento de Ciências da Vida da Faculdade de Ciências da Universidade de Coimbra.
The evolutionary mechanism behind flowers and its pollinators is generally understood to be a gradual co-adaptive process where the plant specializes to its most efficient pollinator, which exerts selective pressures on specific traits, driving floral evolution. Still, most flowering plants in nature are visited by a wide array of pollinator species, i.e. are generalist plants. However, the role of pollinators as significant drivers of floral evolution in generalist plants has been questioned due to the potential conflicting selection regimes exerted by different pollinators. Taking this into account, using a combination of observation and manipulative experiments, we assessed pollinator preference in a natural contact zone where the generalist rayed species Anacyclus clavatus and the rayless A. valentinus co-exist and hybridize, forming intermediate phenotypes. These contact areas show a remarkably high phenotypic variation, with the intermediate phenotype bridging both phenotypes and forming an exceptional micro-evolutive framework to explore how generalist pollinators could be driving the evolution of floral phenotypes. We found that the production of rays influenced the probability of being visited by specific insect groups, in particular by Dipteran groups; whereas bees showed no preference for rayed phenotypes and their visitation patterns were mainly driven by the number of capitula simultaneously blooming in the plant. In addition, we found support for the importance of the neighbours’ phenotype when assessing pollinator preference on a focal individual. Rayed plants benefited from having other conspicuous neighbours, whereas rayless and intermediate phenotypes significantly competed for pollinators. In conclusion, all these differential behavioural patterns of floral visitors might affect gene flow within the hybrid zone between A. clavatus and A. valentinus influencing the degree of reproductive isolation and floral evolution between both species.
Os mecanismos evolutivos que atuam nas flores e respetivos polinizadores são normalmente descritos como processos de co-adaptação gradual onde a planta se especializa no seu polinizador mais eficiente, que por sua vez, exerce pressões evolutivas em características específicas e dessa forma guia a evolução da flor. Ainda assim, a maior parte das plantas com flor são polinizadas por um leque diversificado de espécies de polinizadores, denominando-se assim plantas generalistas. No entanto, em plantas generalistas o papel dos polinizadores na evolução floral tem sido questionado devido a potenciais conflitos na selecção exercida pelas diferentes espécies de polinizadores que visitam a flor. Tendo isto em conta, abordagens observacionais e manipulativas foram utilizadas para avaliar as preferências dos polinizadores numa zona de contacto onde as espécies generalistas Anacyclus clavatus (com lígulas) e a espécie A. valentinus (sem lígulas) coexistem e hibridizam, formando fenótipos intermédios. Estas áreas possuem uma variação fenotípica notável, com o fenótipo intermédio a representar o cruzamento entre as duas espécies, garantindo um cenário microevolutivo excecional para estudar de que forma os polinizadores conduzem a evolução fenotípica em espécies generalistas. Os resultados obtidos revelaram que a produção de lígulas influenciou a probabilidade das plantas serem visitadas por grupos específicos de insetos, em particular por dípteros; por sua vez, as abelhas não revelaram preferências por um fenótipo em particular, preferindo maioritariamente plantas com um elevado número de capítulos em flor. Além disto, os nossos resultados evidenciaram também que a composição fenotípica da vizinhança poderá desempenhar um papel importante na atração de uma planta focal específica; em particular, plantas liguladas beneficiaram em ter outros vizinhos com lígulas, enquanto que os fenótipos sem lígulas e intermédios competiram significativamente por polinizadores. Os diferentes padrões de comportamento diferentes por parte dos visitantes florais observados neste estudo podem afetar o fluxo genético na zona híbrida entre A. clavatus e A. valentinus, influenciando o grau de isolamento reprodutivo e evolução floral entre as duas espécies.

Abstract Invasive plants often occur at high densities and tend to be highly generalist in their interactions with herbivores, pathogens, mycorrhiza, endophytes and pollinators. These characteristics mean that invasive plants should frequently participate in diverse indirect biotic interactions with the surrounding community, mediated by their direct interaction partners (e.g. antagonists and mutualists). Indirect interactions play an important role in many ecological processes, yet we still lack a systematic understanding of the circumstances under which they influence the success and impacts of invasive species. In this chapter, I first describe several of the indirect interaction pathways that are commonly encountered in invasion biology and review their contribution to the impacts of plant invasions on co-occurring species. The literature review revealed that there are now many case studies describing various indirect impacts of invasive plants. However, identical interaction motifs (e.g. plant-enemy-plant, plant-mutualist-plant) can bring about several possible outcomes, depending upon each species' provenance, relative abundances and interaction strengths, abiotic resource availability, spatial and temporal scale and the influence of other species. Moreover, knowledge gaps identified include a lack of studies of indirect facilitation outside of plant-pollinator systems, limited consideration of indirect invader impacts on other non-native species, and the scarcity of generalizable results to date. Second, I integrate the literature with some trending research areas in invasion biology (interaction networks, biogeography, invasion dynamics) and identify some potential future research directions. Finally, I discuss how knowledge about indirect biotic interactions could be incorporated into the management of invasive plants.

Survival and reproduction of several wild plants and crops is mostly by insects pollinator, their recognition and importance have been increased in this climatic changing scenario, which affects the various aspects of their life cycle. According to an estimate, approximately 30,000 species of bees are known in entomology, and about 190 species of bees have been reported to be associated with pollination. There can be an established link between seed production and pollinator diversity, for the plants with a generalist pollination system. The increasing of human habitation affects insect pollinators in various ways, i.e. of habitat destruction, results in low availability of food sources, nesting, oviposition, resting, and mating sites. Pollinator availability restraints the geographical distribution of plant species, i.e. to develop an ecological niche of certain plant species. Failure of pollinator- plant interaction mutualism results in lower seed production and sometimes extirpation of plant population has been recorded. The declining pollinators’ population strengthens existing plant-pollinator interaction or allows new pant pollinator interaction to form. Maintaining the commercial and wild pollinator populations and preventing future shortages of pollination services, therefore, is extremely significant.

Population history and colonization dynamics in the Mediterranean mosaic landscape have closely interacted with natural selection to shape patterns of variation in mating systems and the evolution of floral traits and polymorphisms. Floral phenology is closely adapted to avoid the summer drought regime in many plant communities. Many plant species have generalist pollination systems with a complex of tight and loose interactions that affect the evolution of reproductive traits. Floral traits have clearly accompanied diversification in many groups of plants. The evolution of interactions of plants with their pollinators at the landscape and biogeographic scales illustrates intriguing examples of ongoing evolution.

Invasive plants often occur at high densities and tend to be highly generalist in their interactions with herbivores, pathogens, mycorrhiza, endophytes and pollinators. These characteristics mean that invasive plants should frequently participate in diverse indirect biotic interactions with the surrounding community, mediated by their direct interaction partners (e.g. antagonists and mutualists). Indirect interactions play an important role in many ecological processes, yet we still lack a systematic understanding of the circumstances under which they influence the success and impacts of invasive species. In this chapter, I first describe several of the indirect interaction pathways that are commonly encountered in invasion biology and review their contribution to the impacts of plant invasions on co-occurring species. The literature review revealed that there are now many case studies describing various indirect impacts of invasive plants. However, identical interaction motifs (e.g. plant-enemy-plant, plant-mutualist-plant) can bring about several possible outcomes, depending upon each species' provenance, relative abundances and interaction strengths, abiotic resource availability, spatial and temporal scale and the influence of other species. Moreover, knowledge gaps identified include a lack of studies of indirect facilitation outside of plant-pollinator systems, limited consideration of indirect invader impacts on other non-native species, and the scarcity of generalizable results to date. Second, I integrate the literature with some trending research areas in invasion biology (interaction networks, biogeography, invasion dynamics) and identify some potential future research directions. Finally, I discuss how knowledge about indirect biotic interactions could be incorporated into the management of invasive plants.

This chapter examines the core issues raised by critiques of pollination syndromes and alternative approaches to characterizing pollination in communities, in the context of floral and pollinator specialization. It first considers theoretical arguments against syndromes and specialization before discussing practical evidence against syndromes, along with proposed alternative approaches, focusing in particular on ecological models of specialization and generalization. It then explores some problems with pollination webs, several key issues relating to specialization and generalization, and selection for specialization in flower–pollinator interactions. It also compares patterns of generalization and specialization in different ecosystems and asks whether generalization can be reversed. The chapter concludes with some reflections on the debate over generalization, specialization, and pollination syndromes.

As a way of remedying the wider absence of computer-animated film acting within scholarship on film and animated performance, this chapter makes a significant assertion that, in its production, the computer-animated film genre actually cross-pollinates stop-frame techniques with those associated with marionette theatre as part of its style of performance. In the workable geometry of its virtual bodies (skeletal structure, anatomical coherency, joint segmentation and armature), computer-animated films evoke the wealth of string marionettes (as well as rod or hand puppets) moved within a live performance setting. Such puppet-like forms of acting holds the computer-animated film distinct from performances in popular Hollywood cinema achieved through stop-motion frame-by-frame techniques and traditional hand-drawn methods. However, this analysis not only supports the central concept that puppetry has become a more significant concern of the computer-animated film than in other animated media, but also provides a counter-narrative to scholarship that affords generality to motion-capture as the dominant mode of cyber or virtual puppetry. Puppetry can be understood, I argue, as an altogether more inclusive category, and this chapter promotes puppetry as opening up performance in computer-animated films and revealing the sliding scale of puppet processes involved in its creation of acting.