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.

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Thesis (PhD)--Stellenbosch University, 2011. / ENGLISH ABSTRACT: Both taxonomic diversity and diversity of species interaction networks are experiencing declines as a result of agricultural intensification at habitat and landscape scales. Reversing this trend is a key conservation issue, particularly for important functional groups such as flower-visiting insects and the networks within which they interact. This is of great concern in regions of high conservation priority such as the Cape Floristic Region (CFR), known for its high level of floral and faunal endemism and exceptional species turnover. Holistic approach to conservation in agricultural landscapes involves both preservation of natural land and wildlife friendly management of the farm land to achieve conservation targets. The value of these extensive management approaches is yet to be fully assessed, especially in perennial systems such as vineyards. I examined here the effects of vineyard management and landscape context on species richness and abundance of flower-visiting insects and their species interaction networks. Possible taxon specific effects were verified. I also investigated whether vineyards under organic and conventional management homogenized insect-flower interaction networks and whether vineyards with different management practices vary in patterns of species turnover. I sampled flower-visiting insects and their interactions in organic and conventional vineyards, and in natural reference sites. Inclusion of natural reference sites enabled me to make management recommendations for patches of natural vegetation in CFR agricultural landscape. Statistical models showed taxon-specific benefit of organic farm management, and of landscape (distance to natural habitat). There was benefit to monkey beetles (Scarabaeidae) but not to bees (Apidae). Organic vineyards had a higher number of insect-flower interactions than conventional ones, but vineyards under the two types of management were similar in terms of other important network indices. However, networks of the vineyards were more nested than the natural sites, indicating that they may be potentially more stable to perturbation and random extinctions. Multivariate dispersion tests revealed insect-flower interaction networks were not homogenized by both organic and conventional vineyards across the landscapes. I also found, through additive partitioning, that organic and conventional vineyards were similar in terms of species turnover of bees and flowering plants. The findings of this study provide heuristic value to current debates on the value of vineyard habitats for insect conservation. Both organic and conventional vineyards that promote sustainable management of the non-crop floral vegetation between vineyard rows are potential solutions for conservation of flower-visiting insects and their interactions. Also, attention has to be paid to the quality and connectivity of the natural habitat patches that are within CFR agricultural landscape. Site specific management and assessment of the value of these landscape elements is important. Management approaches such as carefully controlled burning may be beneficial, as the CFR natural vegetation is a fire-driven community. / AFRIKAANSE OPSOMMING: Taksonomiese diversiteit en diversiteit van spesies-interaksie netwerke ondervind beide afnames as gevolg van landboukundige intensifikasie op habitat en landskap skaal. Om die neiging terug te swaai, is ’n sleutel bewaringsaangeleentheid, veral vir belangrike funksionele groepe soos blom-besoekende insekte en die netwerke waarbinne hulle op mekaar inwerk. Dit is van groot kommer in streke met hoë bewaringsprioriteite soos in die Kaapse Floristiese Streek (KFS), bekend vir sy hoë vlak van plant- en dierendemisme en buitengewone spesies kentering. ’n Holistiese benadering tot bewaring van landboukundige landskappe behels beide die bewaring van natuurlike land en natuurlewe-vriendelike bestuur van die plaasgrond om bewaringsdoelwitte te bereik. Die waarde van hierdie ekstensiewe bestuursbenaderings moet nog volledig bepaal word, veral in meerjarige sisteme soos wingerde. Ek het die uitwerkings van wingerdbestuur en landskapsamehang op spesiesrykheid en volopheid van blombesoekende insekte en hulle spesies interaksie netwerke ondersoek. Moontlike takson-spesifieke uitwerkings is nagegaan. Ek het ook ondersoek ingestel of wingerde onder organiese en gebruiklike bestuur ooreenstemmende insek-blom interaksie netwerke met wingerde met verskillende bestuurspraktyke in patroon van spesies kentering gewissel het. Ek het blom-besoekende insekte en hulle interaksies in organiese en konvensionele wingerde, asook in natuurlike verwysingsgebiede gemonster. Insluiting van natuurlike verwysingsgebiede het my in staat gestel om bestuursvoorstelle vir gebiede van natuurlike plantegroei in KFS landboulandskappe voor te stel. Statistiese modelle toon takson-spesifieke voordeel van organiese plaasbestuur en van die landskap (afstand van natuurlike habitat) self. Daar was voordeel vir bobbejaankewers (Scarabaeidae), maar nie vir bye (Apidae) nie. Organiese wingerde het ’n groter getal insek-blom interaksies as konvensionele wingerde gehad, maar wingerde onder beide tipes van bestuur was soortgelyk in terme van ander belangrike netwerk aanduiders. Netwerke van wingerde was egter meer geklomp dan natuurlike gebiede wat aandui dat hulle potensieel meer stabiel betreffende versteuring en lukrake uitsterwings is. Multivariate verspreidingstoetse het aangetoon dat insek-blom interaksie netwerke by beide organiese en konvensionele wingerde oor landskappe nie eenvormig was nie. Ek het ook bevind deur aanvullende verdeling dat organiese en konvensionele wingerde gelykwaardig was in terme van spesies kentering van bye en blomplante. Die bevindings van hierdie studie verskaf heuristise waarde tot huidige debatte oor die waarde van wingerdhabitatte vir insekbewaring. Beide organies en konvensionele wingerde wat volhoubare bestuur van die nie-gewas plantegroei binne wingerdrye bevorder, is moontlike oplossings vir die bewaring van blom-besoekende insekte en hulle wisselwerkings. Bykomend moet aandag gegee word aan die kwaliteit en verbindings van en tussen natuurlike habitat gebiede binne die KFS landboulandskap. Plekspesifieke (plaaslike) bestuur en bepaling van die waarde van hierdie landskapelemente is belangrik. Bestuursbenaderings, soos noukeurig beheerde brand, mag voordelig wees aangesien die KFS natuurlike plantegroei ’n vuurgedrewe gemeenskap is.