Insect pollination of oilseed rape

Ecological intensification of agriculture is suggested as a way to reach higher crop yields without increasing inputs that may degrade the environment. Increased insect pollination in crops, such as oilseed rape, Brassica napus, has been suggested to increase yields, but is rarely integrated in crop management. To determine the value of enhanced crop pollination as a means of ecological intensification, reliable estimates of how yield is affected by insect pollination are needed. Further, little is known about interactions between insect pollination and other crop management factors such as cultivar, nitrogen fertilization, pest control, and irrigation. Finally, it needs to be assessed how increasing crop pollination by adding honey bees to crops impact the wild fauna of flower-visiting insects. I addressed these issues in two sets of experiments. First, I performed a landscape-scale experiment with replicated whole fields of winter oilseed rape. I manipulated the pollinator community by adding honey bee hives or controlled the surroundings for absence of honey bees. I chose fields such that they were embedded in either intensive-ly cropped landscapes or heterogeneous landscapes with more semi-natural pastures, expecting greater diversity of wild pollinators in the latter. In two cultivar types, I examined how honey bee addition affected crop yield and the wild pollinator community. Second, I performed two field plot experiments, in winter and in spring oilseed rape, to assess how insect pollination and the crop management factors cultivar, nitrogen fertili-sation, pest control, and irrigation interactively shaped crop yield. Insect pollination increased winter oilseed rape yield, but only in open-pollinated cultivars. Cultivars of open-pollinated type gave higher yields than cultivars of hybrid type. Thus, phasing out open-pollinated cultivars from the market emerges a missed opportunity for increased yields. I show that the crop’s access to water, nitrogen, and herbivory affect pollinator behaviour and potentially crop pollination. Interestingly, insect pollination tended to increase yields when no nitrogen was applied, indicating higher nutrient use efficiency in plants with access to insect pollination. Adding managed honey bee hives had nega-tive effects on the densities of wild flying- and flower-visiting insects, with potential negative effects on crop pollination and biodiversity conservation. Overall, my thesis demonstrates the importance of including both agronomic and environmental perspectives when developing crop production systems that are productive and sustainable

Legacy of landscape crop diversity enhances carabid beetle species richness and promotes granivores

It is well understood that agricultural expansion and associated loss of semi-natural habitat in the landscape are major drivers for the marked decline in biodiversity. While conserving remaining semi-natural habitat patches is essential to reverse ongoing biodiversity declines, increasing focus has also been put on diversifying cropland itself by increasing landscape crop diversity as a measure of compositional heterogeneity, and reducing field sizes as a measure of configurational heterogeneity. Both these cropland diversification approaches have shown promise to enhance biodiversity in the year of sampling, but it is unknown whether legacies of crop diversity in the landscape promote biodiversity by building up arthropod communities over time. We selected 14 faba bean fields in landscapes dominated by cropland. The fields were chosen along three gradients: landscape crop di-versity of the year of sampling (2017), landscape crop diversity of the previous year (2016) and mean field size in landscapes. Using pitfall traps, we show that the carabid beetle species richness is higher in landscapes with higher crop diversity in the previous year. Especially, granivorous carabid beetles benefitted from legacies of crop diversity. Rove beetles were more abundant and genus rich in landscapes with larger field sizes, while spiders were not responding to any of the landscape variables. A diversity of crops in the landscape and their associated weed communities could provide more diverse food resources and shelter habitats, which build populations of carabid beetle species over time. There is a need to explore the effects of agri-environmental schemes across multiple years to better understand legacy effects, and to structure sustainable agricultural landscapes

Landscape crop diversity and semi-natural habitat affect crop pollinators, pollination benefit and yield

Agricultural intensification has led to the conversion of natural habitats into agricultural fields, increased field sizes and simplified crop rotations. The resulting homogenisation of the landscape has led to a decline in bees, which provide an essential ecosystem service to agriculture. It has been suggested that an increase in landscape crop diversity supports higher biodiversity by providing more diverse and continuous resources without taking land out of agricultural production. We selected 14 faba bean (Vicia faba minor L.) fields in southern Sweden along uncorrelated gradients of landscape crop diversity and proportion of semi-natural habitat within 1.5 km radii surrounding focal fields. Pollinator surveys and pollinator exclusion experiments were conducted to assess whether landscape crop diversity affected pollinator densities, pollinator foraging behaviour (i.e. legitimate flower visitation, nectar robbing or extra-floral nectary visitation), pollination and yield formation. Landscape crop diversity enhanced bumble bee densities. Insect-pollinated faba bean plants produced, on average, 27 % higher bean weight per plant than bagged plants and the insect pollination benefit decreased with increasing semi-natural habitat cover. Bumble bee and honey bee densities, the proportion of nectar robbing bees as well as faba bean yield increased with increasing proportion of semi-natural habitat. Pollinator densities were not the driver of high yields associated with higher proportions of semi-natural habitat because the observed yield increase was unrelated to pollinator densities and driven by bagged plants that were excluded from pollinator visits. Insect pollination, however, clearly decreased the yield gap associated with low proportions of semi natural habitat in the landscape. Our results highlight that agri-environmental policies should promote the retention of existing semi-natural habitats and encourage landscape crop diversity to provide pollinators with sufficient food and nesting resources

Flower strips enhance abundance of bumble bee queens and males in landscapes with few honey bee hives

Wild bee declines in agricultural landscapes have led farmers to supplement crops with honey bees. Simultaneously, environmental subsidy and conservation programmes have incentivized farmers to establish flower strips to support wild and managed pollinators. To find out if flower strips enhance, and competition from honey bees suppresses, wild bees in the landscape and across seasons, we surveyed bumble bee and honey bee abundances in 16 sites in Sweden in summer 2018. The centre of each site (2 km radius) was with or without an annual flower strip, and with or without added honey bee hives. We surveyed bees in each flower strip and in linear habitats in the landscape around each site, such as field edges and road verges. In the following spring, we surveyed bumble bee queen abundance in each site. We show that adding flower strips benefits bumble bee queen abundance the following year, but this effect is diminished if honeybee hives are added. In sites with flower strips, added honey bee hives reduced male bumble bee abundance. Our relatively small flower strip areas bolstered bumble bee population growth across seasons, probably by relieving a resource bottleneck. Adding honey bee hives in combination with flower strips to landscapes with few floral resources should be avoided as it cancelled the positive effect of flower strips

Simple and farmer-friendly bumblebee conservation : Straw bales as nest sites in agricultural landscapes

Many bumblebee species are declining due to a loss of semi-natural habitats in agricultural landscapes resulting in diminished forage and nest sites. Anecdotal experience indicates that bumblebees nest in straw bales, but scientific evidence is lacking. We spent 250 h screening for bumblebee nests in 1255 straw bales and ten straw stacks belonging to 58 farms in two intensively farmed Swedish regions and recorded nests, nest traffic, and straw characteristics. We supplemented the straw screening with screening of control areas, without straw, that were selected in similar environments as the areas with straw. We observed 45 bumblebee nests (including potential nests where a single bumblebee flew in or out of the straw) of eight species/species groups, including one red-listed, in or directly adjacent to the straw at 26 of the farms. Nests were mainly found in partly decayed straw and bales placed together. We found no nests in control areas. Based on our results, we suggest that straw can be used as an easy, cheap and efficient intervention to increase the availability of bumblebee nest sites in agricultural landscapes. Considering the costs and benefits of the alternatives, we conclude that straw addition has advantages over commercial bumblebee colonies for crop pollination purposes and over artificial nest boxes for conservation purposes

Variable pollen viability and effects of pollen load size on components of seed set in cultivars and feral populations of oilseed rape

Pollination success is important for crop yield, but may be cultivar dependent. Less is known about which floral traits influence pollination success. Floral traits, e.g. traits related to attraction and reward, can also contribute to gene flow via pollen, the latter being of particular importance in oilseed rape (Brassica napus) where gene flow occurs between plants of crop, volunteer and feral origin as well as related taxa. We investigated the relationship between pollen load size and seed set in winter oilseed rape. We compared variability in pollen-viability traits, flower production (flowers from the main raceme times number of branches) and seed number and weight per siliqua among cultivars and feral populations (growing outside of agricultural fields) under controlled conditions. Both seed number and weight were saturated at relatively low pollen loads in the tested cultivar. Pollen viability and estimated flower production differed among cultivars, indicating that these traits could contribute to yield variability. Seed weight per siliqua, but not pollen traits or flower production, was lower in ferals compared to cultivars. Thus, while the probability of establishment may be reduced in ferals (due to lower seed weight per siliqua) this will not necessarily impact their contribution to gene flow via pollen. In oilseed rape a relatively low pollen load may be sufficient for full seed set in some cultivars, suggesting less dependence on insect pollination for high yield than generally expected. Our results also showed that previously less investigated floral traits, such as pollen viability, pollen tube growth rate and flower number, can differ between cultivars. Studies of these traits may provide targets for increasing crop yield and provide general knowledge about gene flow between cultivated, feral and related wild populations

Annual flower strips and honeybee hive supplementation differently affect arthropod guilds and ecosystem services in a mass-flowering crop

Intensively managed agricultural landscapes have degraded the provisioning of diverse and continuous forage and shelter habitats for arthropods and weakened the delivery of ecosystem services such as insect crop pollination and biological pest control. In response, farmers are incentivised to sow flower strips along field margins to counteract resource bottlenecks. Yet, it is poorly understood how effective this diversification practice is when combined with the supplementation of honeybee (Apis mellifera L.) hives, which is commonly used to boost insect pollination in flowering crops. Honeybees share floral resources with wild pollinators and natural enemies of pests, which could lead to competition for food resources. We sampled pollinators, natural enemies and their pests as well as estimated the benefit of insect pollination in 17 organic faba bean (Vicia faba minor L.) fields in southern Sweden either with or without sown annual flower strips and with or without added honeybee hives. In fields with flower strips, bumblebee (Bombus spp.) densities were redistributed from field edges to interiors but without affecting their overall densities. Flower strips enhanced silver Y moth (Autographa gamma L.) densities and carabid beetle Shannon diversity along the field edge, and overall spider activity density. The supplementation of honeybee hives enhanced honeybee densities, overall ladybird beetle densities, black bean aphid (Aphis fabae Scop.) densities along field edges, but deterred silver Y moths and pushed bumblebees towards the field interior. Bean mass per plant was higher in insect pollinated plants compared with bagged, self-pollinated plants. This insect pollination benefit was independent of honeybee hive supplementation and the flower strip treatment suggesting that faba bean fields were not deficient in pollinator visits. We conclude that flower strips did not provide sufficient floral resources to increase overall wild pollinator densities in faba bean fields. Yet, annual flower strips attracted and facilitated ground-dwelling predators, especially spiders, to faba bean fields, likely by providing beneficial shelter habitats. It is worth noting that 2018, in which we collected our data, was characterised by late frosts in spring followed by an unusually hot and dry summer. While these unforeseen weather conditions together with a relatively small sample size might limit the generalisation of our results, we argue that conducting experiments under such conditions provide insights into the effectiveness of agri-environmental schemes under climate change, especially considering that such weather conditions are becoming increasingly more frequent

Crop management affects pollinator attractiveness and visitation in oilseed rape

Ecological intensification of agriculture implies managing ecological processes to improve performance of agricultural systems. However, impacts on relevant ecological functions such as insect pollination from other crop management factors are poorly explored. Pest insects and crop resources such as water availability can directly affect crop yields, but it is unknown if there are indirect effects through effects on insect pollination. With a factorial experiment, we examined how irrigation and control of pollen beetles affected crop attractiveness and pollinator visitation in an open-pollinated spring oilseed rape cultivar. We studied how irrigation and pest control modified the production of flowers and nectar in oilseed rape, and if this in turn affected the flower-visitation of honey bees and bumble bees. Pest control increased the number of oilseed rape flowers by 69%, and the amount of nectar per flower with 36%, but for the latter only in non-irrigated plots. Furthermore, we found higher pollinator densities in plots with reduced pollen beetle densities. Pest control also reduced the number of non-legitimate flower visits, suggesting higher pollination efficiency in plots with reduced pollen beetle densities. We show that crop management affects the value of mass-flowering crops as a resource for pollinating insects. Development of pest control tools that are harmless to pollinators could increase the value of flowering crops as food resources for pollinating insects

Data from: Experimental evidence that honeybees depress wild insect densities in a flowering crop

While addition of managed honeybees (Apis mellifera) improves pollination of many entomophilous crops, it is unknown if it simultaneously suppresses the densities of wild insects through competition. To investigate this, we added 624 honeybee hives to 23 fields of oilseed rape (Brassica napus L.) over 2 years and made sure that the areas around 21 other fields were free from honeybee hives. We demonstrate that honeybee addition depresses the densities of wild insects (bumblebees, solitary bees, hoverflies, marchflies, other flies, and other flying and flower-visiting insects) even in a massive flower resource such as oilseed rape. The effect was independent of the complexity of the surrounding landscape, but increased with the size of the crop field, which suggests that the effect was caused by spatial displacement of wild insects. Our results have potential implications both for the pollination of crops (if displacement of wild pollinators offsets benefits achieved by adding honeybees) and for conservation of wild insects (if displacement results in negative fitness consequences)

Data from: Experimental evidence that honeybees depress wild insect densities in a flowering crop

While addition of managed honeybees (Apis mellifera) improves pollination of many entomophilous crops, it is unknown if it simultaneously suppresses the densities of wild insects through competition. To investigate this, we added 624 honeybee hives to 23 fields of oilseed rape (Brassica napus L.) over 2 years and made sure that the areas around 21 other fields were free from honeybee hives. We demonstrate that honeybee addition depresses the densities of wild insects (bumblebees, solitary bees, hoverflies, marchflies, other flies, and other flying and flower-visiting insects) even in a massive flower resource such as oilseed rape. The effect was independent of the complexity of the surrounding landscape, but increased with the size of the crop field, which suggests that the effect was caused by spatial displacement of wild insects. Our results have potential implications both for the pollination of crops (if displacement of wild pollinators offsets benefits achieved by adding honeybees) and for conservation of wild insects (if displacement results in negative fitness consequences)