Food for pollinators: quantifying the nectar and pollen resources of urban flower meadows

Planted meadows are increasingly used to improve the biodiversity and aesthetic amenity value of urban areas. Although many ‘pollinator-friendly’ seed mixes are available, the floral resources these provide to flower-visiting insects, and how these change through time, are largely unknown. Such data are necessary to compare the resources provided by alternative meadow seed mixes to each other and to other flowering habitats. We used quantitative surveys of over 2 million flowers to estimate the nectar and pollen resources offered by two exemplar commercial seed mixes (one annual, one perennial) and associated weeds grown as 300m2 meadows across four UK cities, sampled at six time points between May and September 2013. Nectar sugar and pollen rewards per flower varied widely across 65 species surveyed, with native British weed species (including dandelion, Taraxacum agg.) contributing the top five nectar producers and two of the top ten pollen producers. Seed mix species yielding the highest rewards per flower included Leontodon hispidus, Centaurea cyanus and C. nigra for nectar, and Papaver rhoeas, Eschscholzia californica and Malva moschata for pollen. Perennial meadows produced up to 20x more nectar and up to 6x more pollen than annual meadows, which in turn produced far more than amenity grassland controls. Perennial meadows produced resources earlier in the year than annual meadows, but both seed mixes delivered very low resource levels early in the year and these were provided almost entirely by native weeds. Pollen volume per flower is well predicted statistically by floral morphology, and nectar sugar mass and pollen volume per unit area are correlated with flower counts, raising the possibility that resource levels can be estimated for species or habitats where they cannot be measured directly. Our approach does not incorporate resource quality information (for example, pollen protein or essential amino acid content), but can easily do so when suitable data exist. Our approach should inform the design of new seed mixes to ensure continuity in floral resource availability throughout the year, and to identify suitable species to fill resource gaps in established mixes

The influence of tannins on the extrafloral nectar characteristics and insect mutualists of Vicia faba L.

The main objective of this research project was to investigate the influence of tannins on the extrafloral nectar characteristics and insect mutualists of Vicia faba L. Tannin-free cultivars of V. faba have become increasingly popular in Western Canada due to the greater digestibility of their protein by monogastrics; however, the effect of their lack of tannins on mutualistic insects is unknown. Tannin-rich cultivars of V. faba produce characteristic dark spots on the flowers’ wing petals, and on the stipular extrafloral nectaries (EFNs), which are often used by insects to help locate nectaries. Tannin-free V. faba cultivars lack these nectar guides and spots though, and may be unable to attract as many beneficial insects to the EFNs for herbivore-control purposes, and to the flowers for pollination. Accordingly, this study investigated two tannin-rich (Fatima, SSNS-1) and two tannin-free cultivars (Snowbird, Snowdrop). Extrafloral nectar characteristics were also examined, as the production of tannins can be metabolically expensive, and could come at the cost of extrafloral nectar secretion. Tannin-free cultivars are therefore expected to attract fewer beneficial insects due to their unmarked flowers and EFNs, unless the lack of tannins corresponds with an increased production of nectar or nectar sugars. To examine the effects of tannins on the insect mutualists of V. faba, surveys of insect visitors to the EFNs and flowers were conducted throughout the summers of 2013 and 2014. The vast majority of insect visitors to EFNs were ants (Formicidae), followed by ladybird beetles (Coccinellidae), flies of Camptoprosopella borealis Shewell (Lauxaniidae), and predatory (Vespidae) and parasitoid (Ichneumonidae) wasps, whereas the bees Apis mellifera L. and Bombus nevadensis Cresson were the most common visitors to the flowers. The cultivars which those species were present or absent at during surveys were analyzed using generalized mixed models. The results did not support any consistent differences in insect visitors to plants with tannins, compared to those without, suggesting that the marked difference in the visibility of EFNs on tannin-rich cultivars is not essential for EFN recognition by many insect species. Furthermore, insect visits to EFNs occurred at a highly conserved relative location along the stem, due to a probable increase in nectar production a short distance from the shoot apex. For future reference, stipules at this node were termed the Primary Active EFNs. Extrafloral nectar characteristics were studied in a growth chamber through a combination of nectar sampling by microcapillaries and refractometer measurements to examine nectar volume and sugar concentration, respectively, as well as high performance liquid chromatography to measure the proportion of each of the nectar sugars present. On average, the extrafloral nectar per stipule ± S.E.M had a volume of 0.363 ± 0.021 µL, a nectar sugar concentration (by weight) of 32.5 ± 1.3 %, a nectar sugar mass of 137.6 ± 10.0 µg, and a sugar composition of 54.4 ± 1.0% glucose, 31.1 ± 1.0% fructose, and 14.5 ± 1.0% sucrose. Although extrafloral nectar characteristics varied between cultivars, the variability did not appear dependent on tannin presence or absence, nor did it appear to influence the presence or absence of the abundant ant species Formica neoclara Emery, F. podzolica Francoeur, and Lasius pallitarsis (Provancher), at different cultivars in the field. The increased digestibility provided by tannin-free cultivars of V. faba to monogastrics such as chickens, therefore, does not appear to come at the cost of reduced visitation to the plants by ants and other beneficial insects

On the floral rewards and flower-visitor assemblages of annual urban flower meadow seed mixes

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

Turnover in floral composition explains species diversity and temporal stability in the nectar supply of urban residential gardens

Residential gardens are a valuable habitat for insect pollinators worldwide, but differences in individual gardening practices substantially affect their floral composition. It is important to understand how the floral resource supply of gardens varies in both space and time so we can develop evidence‐based management recommendations to support pollinator conservation in towns and cities. We surveyed 59 residential gardens in the city of Bristol, UK, at monthly intervals from March to October. For each of 472 garden surveys, we combined floral abundances with nectar sugar data to quantify the nectar production of each garden, investigating the magnitude, temporal stability, and diversity and composition of garden nectar supplies. We found that individual gardens differ markedly in the quantity of nectar sugar they supply (from 2 to 1,662 g), and nectar production is higher in more affluent neighbourhoods, but not in larger gardens. Nectar supply peaks in July (mid‐summer), when more plant taxa are in flower, but temporal patterns vary among individual gardens. At larger spatial scales, temporal variability averages out through the portfolio effect, meaning insect pollinators foraging across many gardens in urban landscapes have access to a relatively stable and continuous supply of nectar through the year. Turnover in species composition among gardens leads to an extremely high overall plant richness, with 636 taxa recorded flowering. The nectar supply is dominated by non‐natives, which provide 91% of all nectar sugar, while shrubs are the main plant life form contributing to nectar production (58%). Two‐thirds of nectar sugar is only available to relatively specialised pollinators, leaving just one‐third that is accessible to all. Synthesis and applications. By measuring nectar supply in residential gardens, our study demonstrates that pollinator‐friendly management, affecting garden quality, is more important than the size of a garden, giving every gardener an opportunity to contribute to pollinator conservation in urban areas. For gardeners interested in increasing the value of their land to foraging pollinators, we recommend planting nectar‐rich shrubs with complementary flowering periods and prioritising flowers with an open structure in late summer and autumn

A dataset of nectar sugar production for flowering plants found in urban green spaces

Nectar and pollen are floral resources that provide food for insect pollinators, so quantifying their supplies can help us to understand and mitigate pollinator declines. However, most existing datasets of floral resource measurements focus on native plants found in rural landscapes, so cannot be used effectively for estimating supplies in urban green spaces, where non-native ornamental plants often predominate. We sampled floral nectar sugar in 225 plant taxa found in UK residential gardens and other urban green spaces, focussing on the most common species. The vast majority (94%) of our sampled taxa are non-native, filling an important research gap and ensuring these data are also relevant outside of the United Kingdom. Our dataset includes values of daily nectar sugar production for all 225 taxa and nectar sugar concentration for around half (102) of those sampled. Nectar extraction was conducted according to published methods, ensuring our values can be combined with other datasets. We anticipate that the two main uses of these data are (1) to estimate the nectar production of habitats and landscapes and (2) to identify high-nectar plants of conservation importance. To increase the utility of our data, we provide guidance for scaling nectar values up from single flowers to floral units, as is commonly done in field studies

Nectar and pollen production of Helianthus tuberosus L. – an exotic plant with invasiveness potential

In Central Europe, Helianthus tuberosus L. is a late summer/autumn bloomer (August/November). The disc florets produce both nectar and pollen. Floral reward is available in male-phase flowers (pollen and nectar) and in female-phase flowers (nectar). The floral reward is attractive to a variety of insect visitors (honey bees, wasps, flies and butterflies). The season of blooming as well as the total sugar yield (25.4 – 47.4 kg ha–1) and pollen yield (57.8 – 212.7 kg ha–1) indicate that H. tuberosus is important in the enhancement of food resources for pollinators. The generative reproduction in H. tuberosus is impaired (the species does not set seeds/fruits). However, due to its attractiveness for a variety of pollinators in both rural and urban areas, the spread of H. tuberosus should be monitored. Moreover, its propagation needs to be attended with restrictions

Structure of floral nectaries in Aesculus hippocastanum L.

Representatives of the family Sapindaceae exhibit high morphological diversity of the nectary structure. The present paper shows for the first time the results of micromorphological, anatomical, and ultrastructural analyses of floral nectaries in Aesculus hippocastanum. We have also described the forage and signal attractants of these flowers, which are important for the ecology of pollination. Using light, fluorescence, and electron microscopy, we demonstrated that the A. hippocastanum nectary forming a lobed disc is histologically differentiated into the epidermis with stomata, nectariferous parenchyma, subglandular parenchyma, and vascular bundles reaching the basal part of the nectariferous parenchyma. The use of histochemical assays revealed the presence of insoluble polysaccharides, lipids, terpenoids, and polyphenols including coumarins in the nectary tissues. Nectar is exuded onto the nectary surface via stomata and the permeable cuticle. As indicated by the observation of the ultrastructure of the nectary cells, transport of pre-nectar into parenchymal cells may proceed via the symplast and apoplast. We have also demonstrated that nectar transfer outside the protoplasts of parenchymal cells has a character of granulocrine secretion. A. hippocastanum flowers produce nectar abundantly; one flower secreted on average 2.64 mg of nectar and the concentration of sugars in the nectar was 33%

Oilseed rape (Brassica napus) as a resource for farmland insect pollinators: quantifying floral traits in conventional varieties and breeding systems

This is the final version of the article. Available from the publisher via the DOI in this record.Oilseed rape (OSR; Brassica napus L.) is a major crop in temperate regions and provides an important source ofnutrition to many of the yield-enhancing insect flower visitors that consume floral nectar. The manipulation ofmechanisms that control various crop plant traits for the benefit of pollinators has been suggested in the bid toincrease food security, but little is known about inherent floral trait expression in contemporary OSR varieties orthe breeding systems used in OSR breeding programmes. We studied a range of floral traits in glasshouse-grown, certified conventional varieties of winter OSR to test for variation among and within breeding systems.We measured 24-h nectar secretion rate, amount, concentration and ratio of nectar sugars per flower, and sizesand number of flowers produced per plant from 24 varieties of OSR representing open-pollinated (OP), genicmale sterility (GMS) hybrid and cytoplasmic male sterility (CMS) hybrid breeding systems. Sugar concentrationwas consistent among and within the breeding systems; however, GMS hybrids produced more nectar and moresugar per flower than CMS hybrid or OP varieties. With the exception of ratio of fructose/glucose in OP vari-eties, we found that nectar traits were consistent within all the breeding systems. When scaled, GMS hybridsproduced 1.73 times more nectar resource per plant than OP varieties. Nectar production and amount of nectarsugar in OSR plants were independent of number and size of flowers. Our data show that floral traits of glass-house-grown OSR differed among breeding systems, suggesting that manipulation and enhancement of nectarrewards for insect flower visitors, including pollinators, could be included in future OSR breeding programmes.This work was fundedby the BBSRC, including support from an Insect Pollinators Ini-tiative grant awarded to GAW (BB/I000968/1) that was jointlyfunded by the BBSRC, NERC, the Wellcome Trust, Defra, andthe Scottish Government. Support was also received from HighWycombe Beekeepers’ Association. Rothamsted Researchreceives strategic funding from the Biotechnology and BiologicalSciences Research Council (BBSRC) of the UK

PIN6 is required for nectary auxin response and short stamen development

Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/98417/1/tpj12184-sup-0001-FigS1.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/98417/2/tpj12184.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/98417/3/tpj12184-sup-0004-FigS4.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/98417/4/tpj12184-sup-0003-FigS3.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/98417/5/tpj12184-sup-0002-FigS2.pdfhttp://deepblue.lib.umich.edu/bitstream/2027.42/98417/6/tpj12184-sup-0005-FigS5.pd