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Title: On the floral rewards and flower-visitor assemblages of annual urban flower meadow seed mixes
Author: Godfrey, Thomas George
ISNI:       0000 0004 7224 9243
Awarding Body: University of Edinburgh
Current Institution: University of Edinburgh
Date of Award: 2017
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Flower seed mixes are increasingly used to enhance the biodiversity and amenity values of urban green spaces. Urban or “pictorial” flower seed mixes are often used because they are designed using cultivars and non-native species to provide more colourful and longer-lasting flower displays. Although these seed mixes are effective in providing a high density of large colourful flowers, over an extended season, their value for biodiversity, and in particular the floral rewards they provide for flower-visitors, is largely unknown. The overall aim of my thesis was to assess and improve the value of these new urban habitats as forage resources for flower-visiting insects. My approach was to quantify and compare floral reward provision and insect visitation between meadows grown from three exemplar commercial pictorial flower meadow seed mixes (called Marmalade Annual, Short Annual and Cornfield Annual). I also compared these standard commercial mixes with corresponding ‘nectar-enriched’ formulations, which were designed by increasing the proportional seed weight contribution of selected species predicted to produce high quantities of nectar within each mix. To compare floral rewards and visitation between meadows grown from these seed mixes, I set up a field experiment in Sheffield, UK, using a complete randomised block design with six replicate blocks, each with six 25 m2 plots sown with one of the six seed mix treatments. My first objective was to quantify the floral nectar and pollen rewards provided by each flowering species recorded in the meadows (on the scale of a single flower or inflorescence). My second objective was to use these data to quantify the floral rewards provided per unit area by replicate meadows of different seed mix treatments, testing whether enrichment of seed mixes is an effective method of increasing floral nectar sugar rewards. My third objective was to corroborate/correct my morphology-based flower-visitor identifications using DNA barcoding to screen for misidentifications and morphologically cryptic species. I then used these DNA barcode-based identifications to assess whether there are systematic biases in the structure of flower-visitor networks constructed using molecular taxon identifications compared to traditional morphology-based taxon identifications. My fourth objective was to quantify patterns of insect visitation to meadows, testing whether meadows of different seed mix types attract different flower-visitor assemblages. Meadow floral composition surveys revealed that contamination by unintended horticultural species was widespread across replicate seed mix treatments, with contaminants likely germinating from a seed bank laid down during a failed attempt at this experiment the previous year. Contamination particularly affected Marmalade mixes, mainly because the common contaminant species were often also components of the Short and Cornfield mixes. For example, contaminants contributed on average about a third of nectar sugar mass or pollen volume per unit area in Marmalade mix meadows. Hence, contamination fundamentally undermined the internal validity of seed mix treatments, reducing the ability to directly attribute meadow level patterns in floral rewards or flower-visitors to seed mixes. As result, examination of patterns of floral resource provision and insect visitation were more informative at a species scale. In terms of patterns of insect visitation, Centaurea cyanus received 91% of bumblebee visits, 88% of honeybee visits and 29% of hoverfly visits, whilst T. inodorum received 27% of hoverfly visits. Patterns of bumblebee and honeybee visitation indicated preferential visitation to floral units of Centaurea cyanus. Although this species produced high quantities of nectar sugar mass and pollen volume, this did not differentiate it from other Asteraceae, such as Glebionis segetum, Rudbeckia hirta and Coreopsis tinctoria, which all produced high quantities of both floral rewards. Hence, it is likely that floral traits not measured in this study, such as nectar accessibility (‘nectar-holder depth’) or concentration/volume characteristics (which can affect accessibility due to constraints imposed by feeding morphology), drove patterns of preferential visitation in bumblebees and honeybees to C. cyanus. Given that in the absence of contamination there would have been very few bumblebee or honeybee visitors to Marmalade mix meadows, aesthetically designed pictorial meadows can fail to jointly provide benefits for people and some important flower-visiting insect taxa. DNA barcoding did not change specimen identifications for most morphotaxa. However, splitting and/or lumping processes affected almost one third of morphotaxa, with lumping of morphotaxa the most common type of change. This was in part because males and females from sexually dimorphic species were often separated by morphological identification. These DNA barcode-based changes to visitor taxonomy resulted in consistent minor changes in network size and structure across replicate networks. Lumping of morphotaxa decreased taxon richness, reducing the number of unique links and interaction diversity (the effective number of links). Lumping also increased flower-visitor generality, reducing plant vulnerability and increasing overall network connectance. However, taxonomic changes had no effect on interaction evenness or network specialisation. Thus, for this well-studied fauna, DNA barcode-based flower-visitor networks were systematically biased toward fewer taxa and links, with more generalist visitors and specialist plants. Given that many tropical faunas have more species and are less described than in Britain this pattern may not be replicated in other studies. Further studies in contrasting plant-pollinator communities are required before generalisations can be made about systematic biases between networks constructed using morphological versus molecular data. Overall, meadows grown from annual pictorial flower meadow seed mixes provide abundant floral units per unit area of meadow and are a valuable alternative to traditional horticultural flower beds or amenity grasslands in high profile urban contexts. Nevertheless, care must be taken during design of seed mixes and selection of mixes for planting to ensure that species in the mix provide suitable floral resources for an array of flower-visitors, including bees. This would be aided by the integration of informative measures for candidate species of floral rewards or visitor types and visitation rates during seed mix design.
Supervisor: Stone, Graham ; Walsh, Patrick Sponsor: Biotechnology and Biological Sciences Research Council (BBSRC)
Qualification Name: Thesis (Ph.D.) Qualification Level: Doctoral
EThOS ID:  DOI: Not available
Keywords: cultivars ; non-native species ; urban parks ; nectar rewards ; forage resources ; flower-visiting insects ; flower meadow ; pollen rewards ; floral nectar