The Effects of Nectar-Robbing on Fruit Production in Sparattosperma leucanthum (Bignoniaceae)

Many animals behave as robbers or thieves of floral resources, causing damage to floral tissues or consuming resources used to attract pollinators, or producing effects similar to emasculation by reducing the pollen load in the anthers (which generally results in losses in terms of sexual reproduction). The present work examined the direct and indirect impacts caused by nectar-robbing on the reproductive success of Sparattosperma leucanthum. Different manipulations of the flowers were tested to determine if fruit production was influenced by the perforations made in the floral tissues (direct damage), and if there were changes in visitation frequencies or in the behaviors of effective pollinators (indirect damage). Perforations made by nectar robbers did not lower the reproductive success of the plant species studied. The bee Trigona spinipes was the most frequent visitor and caused the largest perforations in the calyx and corolla of S. leucanthum. Additionally, we noted the occurrence of pollen theft by this same bee in flowers that had been protected against nectar-robbing. These results suggest that if S. leucanthum had developed a mechanism of resistance to robbery by T spinipes it would probably have experienced even lower pollination levels as a result of reductions in the quantities of pollen available for transfer by effective pollinators. We were not able to evaluate if nectar depletion through robbery modified the behavior of the effective pollinators (bumblebees of the genus Bombus).

Pollination, Herbivory, and Habitat Fragmentation: Their Effects on the Reproductive Fitness of Angadenia berteroi, a Native Perennial Plant of the South Florida Pine Rocklands

Angadenia berteroi is a tropical perennial subshrub of the pine rocklands with large yellow flowers that set very few fruits. My dissertation seeks to elucidate the factors that affect the reproductive fitness of Angadenia berteroi a native species of the south Florida pine rocklands. I provide novel information on the pollination biology of this native species. I also assess the effects of herbivory on growth and the reproductive success of A. berteroi. Finally, I elucidate how habitat fragmentation and quality are correlated with reproductive fitness of this native perennial plant. Using a novel experimental approach, I determined the most effective pollinator group. I used nylon fishing line of widths corresponding to proboscis diameter of the major groups of visitors to examine pollen removal and deposition. In the field, I estimated visitation frequency and efficacy of each pollinator type. Using potted plants, I exposed flowers to single visit from different types of pollinators to measure fruit set. I performed artificial defoliation with scissors on plants growing in the greenhouse to assess the effects of defoliation before flowering as well as during flowering. Additionally, I used structural equation modelling (SEM) to elucidate how A. berteroi reproductive fitness was affected by habitat fragmentation and quality. My experiments provide evidence that Angadenia berteroi is specialized for bee pollination; though butterflies, skippers and others also visit its flowers, A. berteroi is exclusively pollinated by two native bees of the South Florida pine rocklands . This research also demonstrated that herbivory by the oleander moth may have direct and indirect effects on Angadenia berteroi growth and reproductive success. The SEM results suggested that habitat quality (litter depth and subcanopy cover) may favor reproduction in native species of the South Florida pine rocklands that are properly maintained by periodic fires and exotic control. Insights from this threatened and charismatic species may provide impetus to properly manage remaining pine rocklands in South Florida for this and other endemic understory species.

Pollination, herbivory, and habitat fragmentation: Their effects on the reproductive fitness of Angadenia berteroi, a native perennial plant of the south Florida pine rocklands

Angadenia berteroi is a tropical perennial subshrub of the pine rocklands with large yellow flowers that set very few fruits. My dissertation seeks to elucidate the factors that affect the reproductive fitness of Angadenia berteroi a native species of the south Florida pine rocklands. I provide novel information on the pollination biology of this native species. I also assess the effects of herbivory on growth and the reproductive success of A. berteroi. Finally, I elucidate how habitat fragmentation and quality are correlated with reproductive fitness of this native perennial plant.^ Using a novel experimental approach, I determined the most effective pollinator group. I used nylon fishing line of widths corresponding to proboscis diameter of the major groups of visitors to examine pollen removal and deposition. In the field, I estimated visitation frequency and efficacy of each pollinator type. Using potted plants, I exposed flowers to single visit from different types of pollinators to measure fruit set. I performed artificial defoliation with scissors on plants growing in the greenhouse to assess the effects of defoliation before flowering as well as during flowering. Additionally, I used structural equation modelling (SEM) to elucidate how A. berteroi reproductive fitness was affected by habitat fragmentation and quality. ^ My experiments provide evidence that Angadenia berteroi is specialized for bee pollination; though butterflies, skippers and others also visit its flowers, A. berteroi is exclusively pollinated by two native bees of the South Florida pine rocklands . This research also demonstrated that herbivory by the oleander moth may have direct and indirect effects on Angadenia berteroi growth and reproductive success. The SEM results suggested that habitat quality (litter depth and subcanopy cover) may favor reproduction in native species of the South Florida pine rocklands that are properly maintained by periodic fires and exotic control. Insights from this threatened and charismatic species may provide impetus to properly manage remaining pine rocklands in South Florida for this and other endemic understory species.^

Reproductive Biology of Corymbia citriodora subsp. variegata and Effective Pollination across its Native Range.

The spotted gum species complex represents a group of four eucalypt hardwood taxa that have a native range that spans the east coast of Australia, with a morphological cline from Victoria to northern Queensland. Of this group, Corymbia citriodora subsp. variegata (CCV) is widespread in south-eastern Queensland and northern New South Wales. It is currently the most commonly harvested native hardwood in Queensland. However, little basic knowledge of the reproductive biology of the species is available to inform genetic improvement and resource management programmes. Here we take an integrative approach, using both field and molecular data, to identify ecological factors important to mating patterns in native populations of CCV. Field observation of pollinator visitation and flowering phenology of 20 trees showed that foraging behaviour of pollinator guilds varies depending on flowering phenology and canopy structure. A positive effect of tree mean flowering effort was found on insect visitation, while bat visitation was predicted by tree height and by the number of trees simultaneously bearing flowers. Moreover, introduced honeybees were observed frequently, performing 73% of detected flower visits. Conversely, nectar-feeding birds and mammals were observed sporadically with lorikeets and honeyeaters each contributing to 11% of visits. Fruit bats, represented solely by the grey-headed flying fox, performed less than 2% of visits. Genotyping at six microsatellite markers in 301 seeds from 17 families sampled from four of Queensland's native forests showed that CCV displays a mixed-mating system that is mostly outcrossing (tm = 0.899 ± 0.021). Preferential effective pollination from near-neighbours was detected by means of maximum-likelihood paternity analysis with up to 16% of reproduction events resulting from selfing. Forty to 48% of fertilising pollen was also carried from longer distance (>60 m). Marked differences in foraging behaviour and visitation frequency between observed pollinator guilds suggests that the observed dichotomy of effective pollen movement in spotted gums may be due to frequent visit from introduced honeybees favouring geitonogamy and sporadic visits from honeyeaters and fruit bats resulting in potential long-distance pollinations.

Plant-pollinator biodiversity and pollination services in a complex Mediterranean landscape

Mediterranean landscapes comprise a complex mosaic of different habitats that vary in the diversity of their floral communities, pollinator communities and pollination services. Using the Greek Island of Lesvos as a model system, we assess the biodiversity value of six common habitats and measure ecosystemic 'health' using pollen grain deposition in three core flowering plants as a measure of pollination services. Three fire-driven habitats were assessed: freshly burnt areas, fully regenerated pine forests and intermediate age scrub; in addition we examined oak woodlands, actively managed olive groves and groves that had been abandoned from agriculture. Oak woodlands, pine forests and managed olive groves had the highest diversity of bees. The habitat characteristics responsible for structuring bee communities were: floral diversity, floral abundance, nectar energy availability and the variety of nectar resources present. Pollination services in two of our plant species, which were pollinated by a limited sub-set of the pollinator community, indicated that pollination levels were highest in the burnt and mature pine habitats. The third species, which was open to all flower visitors, indicated that oak woodlands had the highest levels of pollination from generalist species. Pollination was always more effective in managed olive groves than in abandoned groves. However, the two most common species of bee, the honeybee and a bumblebee, were not the primary pollinators within these habitats. We conclude that the three habitats of greatest overall value for plant-pollinator communities and provision of the healthiest pollination services are pine forests, oak woodland and managed olive groves. We indicate how the highest value habitats may be maintained in a complex landscape to safeguard and enhance pollination function within these habitats and potentially in adjoining agricultural areas. (c) 2005 Elsevier Ltd. All rights reserved.

Delivery of crop pollination services is an insufficient argument for wild pollinator conservation

There is compelling evidence that more diverse ecosystems deliver greater benefits to people, and these ecosystem services have become a key argument for biodiversity conservation. However, it is unclear how much biodiversity is needed to deliver ecosystem services in a cost-effective way. Here we show that, while the contribution of wild bees to crop production is significant, service delivery is restricted to a limited subset of all known bee species. Across crops, years and biogeographical regions, crop-visiting wild bee communities are dominated by a small number of common species, and threatened species are rarely observed on crops. Dominant crop pollinators persist under agricultural expansion and many are easily enhanced by simple conservation measures, suggesting that cost-effective management strategies to promote crop pollination should target a different set of species than management strategies to promote threatened bees. Conserving the biological diversity of bees therefore requires more than just ecosystem-service-based arguments.

Apple pollination: demand depends on variety and supply depends on pollinator identity

Insect pollination underpins apple production but the extent to which different pollinator guilds supply this service, particularly across different apple varieties, is unknown. Such information is essential if appropriate orchard management practices are to be targeted and proportional to the potential benefits pollinator species may provide. Here we use a novel combination of pollinator effectiveness assays (floral visit effectiveness), orchard field surveys (flower visitation rate) and pollinator dependence manipulations (pollinator exclusion experiments) to quantify the supply of pollination services provided by four different pollinator guilds to the production of four commercial varieties of apple. We show that not all pollinators are equally effective at pollinating apples, with hoverflies being less effective than solitary bees and bumblebees, and the relative abundance of different pollinator guilds visiting apple flowers of different varieties varies significantly. Based on this, the taxa specific economic benefits to UK apple production have been established. The contribution of insect pollinators to the economic output in all varieties was estimated to be £92.1M across the UK, with contributions varying widely across taxa: solitary bees (£51.4M), honeybees (£21.4M), bumblebees (£18.6M) and hoverflies (£0.7M). This research highlights the differences in the economic benefits of four insect pollinator guilds to four major apple varieties in the UK. This information is essential to underpin appropriate investment in pollination services management and provides a model that can be used in other entomolophilous crops to improve our understanding of crop pollination ecology.

A Meta-Analysis of Predation Risk Effects on Pollinator Behaviour

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Pollinator responses to variation in floral display and flower size in dioecious Sagittaria latifolia (Alismataceae)

In animal-pollinated plants with unisexual flowers, sexual dimorphism in floral traits may be the consequence of pollinator-mediated selection. Experimental investigations of the effects of variation in flower size and floral display on pollinator visitation can provide insights into the evolution of floral dimorphism in dioecious plants. Here, we investigated pollinator responses to experimental arrays of dioecious Sagittaria latifolia in which we manipulated floral display and flower size. We also examined whether there were changes in pollinator visitation with increasing dimorphism in flower size. In S. latifolia, males have larger flowers and smaller floral displays than females. Visitation by pollinators, mainly flies and bees, was more frequent for male than for female inflorescences and increased with increasing flower size, regardless of sex. The number of insect visits per flower decreased with increasing floral display in males but remained constant in females. Greater sexual dimorphism in flower size increased visits to male inflorescences but had no influence on the number of visits to female inflorescences. These results suggest that larger flower sizes would be advantageous to both females and males, and no evidence was found that females suffer from increased flower-size dimorphism. Small daily floral displays may benefit males by allowing extended flowering periods and greater opportunities for effective pollen dispersal.

Support for the predictions of the pollinator-mediated stabilizing selection hypothesis

Aims Floral traits are frequency used in traditional plant systematics because Of their assumed constancy. One potential reason for the apparent constancy of flower size is that effective pollen transfer between flowers depends oil the accuracy of the physical fit between the flower and pollinator. Therefore, dowels are likely to he under stronger stabilizing selection for uniform size than vegetative plant parts. Moreover, as predicted by the pollinator-mediated stabilizing selection (PMSS) hypothesis, all accurate fit between flowers and their pollinators is likely to he more important for specialized pollination systems as found in many species with bilaterally symmetric (zygomorphic) flowers than for species, with radially symmetric (actinomorphic) flowers. Methods In a comparative study of 15 zygomorphic and 13 actinomorphic species ill Switzerland, we tested whether variation in flower size, among and within individuals, is smaller than variation ill leaf size and whether variation in flower size is smaller ill zygomorphic compared to actinomorphic species. Important findings Indeed, variation ill leaf length was significantly larger than variation in flower length and width. Within-individual variation ill flower and leaf sizes did not differ significantly between zygomorphic and actinomorphic species. In line with the predictions of the PMSS, among-individual variation ill flower length and flower width was significantly smaller for zygomorphic species than for actinomorphic species, while the two groups did not differ in leaf length variation. This suggests that plants with zygomorphic flowers have undergone stronger selection for uniform flowers than plants with actinomorphic flowers. This supports that the uniformity of flowers compared to vegetative structures within species, as already observed in traditional plant systematics, is, at least in part, a consequence of the requirement for effective pollination.

Apple Pollination: Demand Depends on Variety and Supply Depends on Pollinator Identity

Insect pollination underpins apple production but the extent to which different pollinator guilds supply this service, particularly across different apple varieties, is unknown. Such information is essential if appropriate orchard management practices are to be targeted and proportional to the potential benefits pollinator species may provide. Here we use a novel combination of pollinator effectiveness assays (floral visit effectiveness), orchard field surveys (flower visitation rate) and pollinator dependence manipulations (pollinator exclusion experiments) to quantify the supply of pollination services provided by four different pollinator guilds to the production of four commercial varieties of apple. We show that not all pollinators are equally effective at pollinating apples, with hoverflies being less effective than solitary bees and bumblebees, and the relative abundance of different pollinator guilds visiting apple flowers of different varieties varies significantly. Based on this, the taxa specific economic benefits to UK apple production have been established. The contribution of insect pollinators to the economic output in all varieties was estimated to be £92.1M across the UK, with contributions varying widely across taxa: solitary bees (£51.4M), honeybees (£21.4M), bumblebees (£18.6M) and hoverflies (£0.7M). This research highlights the differences in the economic benefits of four insect pollinator guilds to four major apple varieties in the UK. This information is essential to underpin appropriate investment in pollination services management and provides a model that can be used in other entomolophilous crops to improve our understanding of crop pollination ecology.

Effective Cell-Centred Time-Domain Maxwell's Equations Numerical Solvers

This research work analyses techniques for implementing a cell-centred finite-volume time-domain (ccFV-TD) computational methodology for the purpose of studying microwave heating. Various state-of-the-art spatial and temporal discretisation methods employed to solve Maxwell's equations on multidimensional structured grid networks are investigated, and the dispersive and dissipative errors inherent in those techniques examined. Both staggered and unstaggered grid approaches are considered. Upwind schemes using a Riemann solver and intensity vector splitting are studied and evaluated. Staggered and unstaggered Leapfrog and Runge-Kutta time integration methods are analysed in terms of phase and amplitude error to identify which method is the most accurate and efficient for simulating microwave heating processes. The implementation and migration of typical electromagnetic boundary conditions. from staggered in space to cell-centred approaches also is deliberated. In particular, an existing perfectly matched layer absorbing boundary methodology is adapted to formulate a new cell-centred boundary implementation for the ccFV-TD solvers. Finally for microwave heating purposes, a comparison of analytical and numerical results for standard case studies in rectangular waveguides allows the accuracy of the developed methods to be assessed.