Role of juvenile hormone biosynthesis in dominance status and reproduction of the bumblebee,Bombus terrestris

Summary In Bombus terrestris females, dominant status is correlated with high levels of juvenile hormone (JH) biosynthesis and rapid oöcyte maturation. Queenright workers, which are inhibited by the dominant queen, complete the cycle of oöcyte maturation while exhibiting a continuous low rate of JH production, but their egglaying is inhibited. Measurements in foundress queens suggest that the low JH titer during oögenesis is probably not responsible for the inhibition of egg-laying. Queenless workers, kept individually, are not activated either for JH production or oöcyte maturation. In groups of three queenless workers, a dominance order becomes established and high rates of JH synthesis are observed in the dominant egg-laying workers, with low rates in subordinated workers. In groups of founder queens, also, a dominance order becomes established and results in a reduced rate of JH production in subordinated females. Correspondence to: F. Couillaud     

Pollinator insects benefit from rotational fallows

The importance of rotational fallows for pollinating insects in intensively cultivated farmland was studied in a three-year field experiment. The focus was on the establishment method (undersowing or seed mixture sown post-tillage), seed mixture (competitive Festuca pratensis-Phleum pratense-Trifolium pratense or less competitive Agrostis capillaris-Festuca ovina mixture) and duration (1 or 2 years) of the fallow. The seed mixture and duration affected the pollinators more than the method of establishment. Pollinator species richness was higher in fallows sown with less competitive than with competitive grasses and it increased in the second year. Stubble fields reached as high pollinator species richness already during the first year as fallows with less competitive grasses did in the second year. Pollinator abundance was highest in those treatments in which species richness of flowering plants was highest. The results suggest that the benefits of rotational fallows for pollinators increase when they are established as stubble or using seeds of less competitive grasses and when the field is fallowed for (at least) two summers.     

Analysis of bumblebee visitation sequences within single bouts: implication of the overstrike effect on short-term memory

Pollinators whose foraging habitats consist of several plant types (species or morph) may continue to choose the plant type last visited because information about the type of plant last visited dominates over all other memory contents, in particular of short-term memory. In this study, I extracted this overstrike effect on the plant choices of pollinators by analyzing patterns of visitation sequences within a single round-trip between the hive and foraging patch (bout). First, I simulated the visitation sequences within single bouts with a model to show how factors, including the bees plant-type preferences, the arrangement of plants and the effect of overstrike on short-term memory, affect visitation sequences. Here, bees are assumed to forage in a patch consisting of two plant types (H and L). The model predicts that only the effect of overstrike on short-term memory causes assorted visitation sequences according to plant type (within-bout flower constancy). That is, if the overstrike-effect on short-term memory is the primary determinant of plant choice, then bees will fly to a type-L plant after visiting a type-L plant even if they predominantly visit type-H plants and vice versa. Next, I investigated individual bumblebees visitation sequences at a patch of artificial inflorescences with a set-up similar to that assumed in the model. Two types of inflorescences were arranged on a Cartesian grid. Assorted visitation sequences according to inflorescence type were observed, depending on the distances among inflorescences. This result supports the hypotheses that bees fly to the same plant type as that last visited because short-term memory is displaced (overstruck) with information about the most recently visited plant type.Communicated by M. Giurfa     

Does interaction between bumblebees (Bombus ignitus) reduce their foraging area?: bee-removal experiments in a net cage

To examine whether the interaction between bumblebees, Bombus ignitus, reduces their foraging area, we conducted bee-removal experiments in a net cage. In the cage, we set potted Salvia farinacea plants, allowed bumblebees to forage freely on those plants, and followed their plant-to-plant movements to identify a bee with a relatively small foraging area. We then removed all the other foraging bees, except for the bee with a small foraging area, and observed the change of the foraging area of the focal bee under conditions of no interaction with other bees. After the removal of the other bees, all five bees tested enlarged their foraging areas, suggesting that the interaction between bees is an important determinant of their foraging areas. The result also means that bumblebees are able to adjust their foraging areas in response to other foragers, indicating the necessity for future studies to clarify what cues bees use to interact with other bees. Moreover, after the removal treatments, all five bees showed temporary increases in the number of flower probes per plant. This can be explained by their optimal foraging according to the old average intake rate for the plant population and by the delayed changes in response to the new high average energy intake rate after the bee-removal treatments.Communicated by M. Giurfa     

The dynamics of social learning in an insect model,the bumblebee (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)

Bumblebees (Bombus terrestris) are attracted to those particular inflorescences where other bees are already foraging, a process known as local enhancement. Here, we use a quantitative analysis of learning in a foraging task to illustrate that this attraction can lead bees to learn more quickly which flower species are rewarding if they forage in the company of experienced conspecifics. This effect can also be elicited by model bees, rather than live demonstrators. We also show that local enhancement in bumblebees most likely reflects a general attraction to conspecifics that is not limited to a foraging context. Based on the widespread occurrence of both local enhancement and associative learning in the invertebrates, we suggest that social influences on learning in this group may be more common than the current literature would suggest and that invertebrates may provide a useful model for understanding how learning processes based on social information evolve.     

Anther-mimicking floral guides exploit a conflict between innate preference and learning in bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)

Bee-pollinated plants are frequently dichogamous: e.g. each flower has a discernable male and female phase, with only the male phase offering a pollen reward. Pollen-collecting bees should therefore discriminate against female-phase flowers to maximise their rate of pollen harvest, but this behaviour would reduce plant fitness due to inferior pollination. Here, we test the hypothesis that flowers use pollen-mimicking floral guides to prevent flower-phase discrimination. Such floral guides resemble pollen in spectral reflection properties and are widespread among angiosperm flowers. In an array of artificial flowers, bumblebees learned less well to discriminate between flower variants simulating different flowering phases when both flower variants carried an additional pollen-yellow guide mark. This effect depended crucially on the pollen-yellow colour of the guide mark and on its spatial position within the artificial flower. We suggest that floral guides evolved to inhibit flower-phase learning in bees by exploiting the innate colour preferences of their pollinators.     

Effects of pollinator density-dependent preferences on field margin visitations in the midst of agricultural monocultures: A modelling approach

Managed field margins offer a means of reducing the impact of agricultural monocultures within intensively managed environments. By providing refuge for wild plants and the pollinators associated with them, field margins can also contribute to enhancing the pollination services within the monoculture. However, the effects of the monoculture on pollinator behaviour need to be carefully considered. It is known that pollinators may show density-dependent preferences such as neophobia (an avoidance of unfamiliar items) when different types of flower are available within their environment, and the dominance of monoculture crops within the environment may consequently have adverse effects upon the preferences shown by pollinators living in the field margins within them. In order to examine how pollinator preferences for wild flowers are affected by monocultures, we modelled the effects of density-dependent preferences, flower densities, and the geometry of field margins within a monoculture landscape using numerical simulations. This was done by considering how the placement of pollinator nests within a simple, spatially explicit landscape consisting of fields of monoculture crops separated by margins containing wild flowers affected the ratio of wild and monoculture crops experienced by the pollinator, given that it could only forage within a limited distance from its nest. Increasing field margin width and decreasing monoculture field width both led to an increase in pollinators visiting wild flowers (which levelled off as width increased). The size of the monoculture fields had little additional effect once they had passed an intermediate width. Increasing wild flower density within the margins led to a shift away from preferring monocultures. When wild flowers were at low densities compared to the monoculture, even the addition of small amounts of extra wild flowers had a large effect in shifting foraging preferences away from the monoculture. The distance which pollinators normally forage over only has an effect upon preferences for wild flowers when the travel distance is small. This suggests that careful consideration of margin design might be extremely important for those species which do not travel far. Innate preferences for density-dependence and particular crop types may also have an effect on preference behaviour. We demonstrate that the way in which resources are presented to indigenous pollinators may be extremely important in influencing where they choose to forage within agricultural landscapes. Careful margin design, as well as increasing the density of wild flowers (such as by enhancing the wild seed bank within the margins), may lead to overall improvements in ecosystem function within intensively farmed monocultures.