Thomas C. Ings

Interspecific competition, whereby individuals of one species are in competition with individuals of another species, occurs naturally throughout invertebrate communities, and in pollinators, it manifests as exploitative or interference competition. Exploitative competition has been suggested as the primary mechanism of competition between managed bees and other pollinators; however, interference competition may also be a significant factor. Apis mellifera Linnaeus, 1758 (honeybee), primarily a managed bee species, where highly abundant, has a negative impact on native non‐ Apis pollinators. It is important to understand the nature of competition between wild and managed species to better conserve wild pollinators. The results presented here add new depth to existing literature about behavioural interactions among honeybees and non‐ Apis pollinators, evidencing interference competition and categorising the incidence of focal behaviours on highly utilised seasonal resources. We observed the behaviours of pollinators foraging on Rubus fruticosus Linnaeus, 1753 (bramble) and Hedera helix Linnaeus, 1753 (ivy), high‐value resources, which are a seasonally important food source for a wide range of pollinators in periods where naturally occurring flowers are scarce. Observations took place across 2 years at UK sites in close proximity to honeybee hives. We documented the outcome of 300 pairs of interactions, amounting to 93 h of observation time. The outcomes were recorded as ‘win’ where one individual gains access to the resource, resulting in a ‘loss’ of forage for the opposite interacting partner, or ‘share’ where two gain access to the resource in parallel with no interaction. Overall, honeybee abundance strongly negatively influenced the abundance of non‐ Apis pollinators. Competitive interactions were more frequent and led to greater than expected ‘wins’ for honeybees. Honeybees were avoided in 28.4% of interactions on bramble and 35.6% on ivy. The honeybee displaced or gave chase when interacting with another pollinator in at least 20% of winning outcomes on both plant species. We conclude that honeybees dominate shared resources by restricting the access of other pollinators to resources and that the importance of interference competition in pollinator communities is being underestimated.

Understanding how ecological communities are structured is a major goal in ecology. Ecological networks representing interaction patterns among species have become a powerful tool to capture the mechanisms underlying plant-animal assemblages. However, these networks largely do not account for inter-individual variability and thus may be limiting our development of a clear mechanistic understanding of community structure. In this study, we develop a new individual-trait based approach to examine the importance of individual plant and pollinator functional size traits (pollinator thorax width and plant nectar holder depth) in mutualistic networks. We performed hierarchical cluster analyses to group interacting individuals into classes, according to their similarity in functional size. We then compared the structure of bee-flower networks where nodes represented either species identity or trait sets. The individual trait-based network was almost twice as nested as its species-based equivalent and it had a more symmetric linkage pattern resulting from of a high degree of size-matching. In conclusion, we show that by constructing individual trait-based networks we can reveal important patterns otherwise difficult to observe in species-based networks and thus improve our understanding of community structure. We therefore recommend using both trait-based and species-based approaches together to develop a clearer understanding of the properties of ecological networks.

A speed-accuracy trade-off (SAT) in behavioural decisions is known to occur in a wide range of vertebrate and invertebrate taxa. Accurate decisions often take longer for a given condition, while fast decisions can be inaccurate in some tasks. Speed-accuracy tactics are known to vary consistently among individuals, and show a degree of flexibility during colour discrimination tasks in bees. Such individual flexibility in speed-accuracy tactics is likely to be advantageous for animals exposed to fluctuating environments, such as changes in predation threat. We therefore test whether individual speed-accuracy tactics are fixed or flexible under different levels of predation threat in a model invertebrate, the bumblebee Bombus terrestris. The flexibility of speed-accuracy tactics in a foraging context was tested in the laboratory using a “meadow” of artificial flowers harbouring “robotic” crab spider predators. We found that while the ranking of bees along the speed and accuracy continuums was consistent across two levels of predation threat, there was some flexibility in the tactics used by individual bees – most bees became less accurate at colour discrimination when exposed to predation threat when flower types were rewarding. The relationship between decision speed and accuracy was influenced by predator detectability and the risk associated with making incorrect choices during the colour discrimination task. Predator crypsis resulted in a breakdown in the relationship between speed and accuracy, especially when making an incorrect floral choice incurred a distasteful quinine punishment. No single speed-accuracy tactic was found to be optimal in terms of foraging efficiency under either predation threat situation. However, bees that made faster decisions achieved higher nectar collection rates in predator free situations, while accurate bees achieved higher foraging rates under predation threat. Our findings show that while individual bees remain relatively consistent in terms of whether they place greater emphasis on speed or accuracy under predation threat, they can respond flexibly to the additional time costs of detecting predators.

Simulated annealing is a powerful stochastic search algorithm for locating a global maximum that is hidden among many poorer local maxima in a search space. It is frequently implemented in computers working on complex optimization problems but until now has not been directly observed in nature as a searching strategy adopted by foraging animals. We analysed high-speed video recordings of the three-dimensional searching flights of bumblebees (Bombus terrestris) made in the presence of large or small artificial flowers within a 0.5 m³ enclosed arena. Analyses of the three-dimensional flight patterns in both conditions reveal signatures of simulated annealing searches. After leaving a flower, bees tend to scan back-and forth past that flower before making prospecting flights (loops), whose length increases over time. The search pattern becomes gradually more expansive and culminates when another rewarding flower is found. Bees then scan back and forth in the vicinity of the newly discovered flower and the process repeats. This looping search pattern, in which flight step lengths are typically power-law distributed, provides a relatively simple yet highly efficient strategy for pollinators such as bees to find best quality resources in complex environments made of multiple ephemeral feeding sites with nutritionally variable rewards.

We analyze 3D flight paths of bumblebees searching for nectar in a laboratory experiment with and without predation risk from artificial spiders. For the flight velocities we find mixed probability distributions reflecting the access to the food sources while the threat posed by the spiders shows up only in the velocity correlations. The bumblebees thus adjust their flight patterns spatially to the environment and temporally to predation risk. Key information on response to environmental changes is contained in temporal correlation functions, as we explain by a simple emergent model.

Foraging bumblebees are normally associated with spring and summer in northern Europe. However, there have been sightings of the bumblebee Bombus terrestris during the warmer winters in recent years in southern England. But what floral resources are they relying upon during winter and how much winter forage can they collect? To test if urban areas in the UK provide a rich foraging niche for bees we set up colonies of B. terrestris in the field during two late winter periods (2005/6 & 2006/7) in London, UK, and measured their foraging performance. Fully automatic radio-frequency identification (RFID) technology was used in 2006/7 to enable us to record the complete foraging activity of individually tagged bees. The number of bumblebees present during winter (October 2007 to March 2008) and the main plants they visited were also recorded during transect walks. Queens and workers were observed throughout the winter, suggesting a second generation of bee colonies active during the winter months. Mass flowering shrubs such as Mahonia spp. were identified as important food resources. The foraging experiments showed that bees active during the winter can attain nectar and pollen foraging rates that match, and even surpass, those recorded during summer. B. terrestris in the UK are now able to utilise a rich winter foraging resource in urban parks and gardens that might at present still be under-exploited, opening up the possibility of further changes in pollinator phenology.