Chez les pollinisateurs sociaux, comme l'abeille domestique (Apis mellifera L.) et le bourdon terrestre (Bombus terrestris L.), mes deux modèles d'étude, différents individus sont spécialisés dans différentes tâches. Il est admis que différents types de comportement de butinage contribuent à une optimisation des performances de la colonie. Actuellement, les populations de pollinisateurs sont exposées à des stress environnementaux, qui sont connus pour perturber le comportement des individus en visant directement leur cognition. Il est ainsi crucial de mieux comprendre comment les colonies d'abeilles et de bourdons maintiennent une activité de butinage efficace, et quels sont les effets de stress environnementaux sur les butineuses. Dans cette thèse, j'ai donc examiné les différentes stratégies de butinage pour différentes sources de nourriture, pollen et nectar, et les variabilités interindividuelles dans le comportement de butinage. Je me suis aussi intéressé à l'impact de stress tels que les pesticides sur l'efficacité de butinage. J'ai utilisé la technologie RFID pour suivre le comportement des abeilles tout au long de leur vie. J'ai trouvé que les colonies d'abeilles et de bourdons reposent sur un petit groupe d'individus très actifs qui fournissent la majorité de la nourriture pour la colonie. Chez les abeilles, ces individus très actifs sont aussi plus efficaces pour collecter nectar et pollen. J'ai aussi identifié l'existence de différentes stratégies pour la collecte de pollen ou de nectar. Ensuite, j'ai pu montrer que les bourdons ont des différences interindividuelles très marquées dans un test de navigation, une tâche cruciale dans le comportement de butinage. Finalement, j'ai testé l'effet néfaste de pesticides sur l'apprentissage visuel chez l'abeille. Cette thèse a pour but de mieux comprendre les causes de vulnérabilité des pollinisateurs aux stress environnementaux. Mes résultats soulignent le besoin de considérer la diversité comportementale comme une adaptation des espèces de pollinisateurs sociaux, mais aussi comme une potentielle cause de vulnérabilité de la colonie vis-à-vis des stress. / In social insects, such as bees, different individuals specialise in the collection of different resources, and it is assumed that natural behavioural variability among foragers contributes to a self-organised optimisation of colony performance. Currently, bee populations are facing an increasing number of environmental stressors, known to disturb the behaviour of individuals, presumably upon their impact on cognitive capacities. Hence it is important to learn more about how stressors impact on individual foraging behaviour to understand how a colony maintains effective nutrition and development. In this thesis in cognitive ecology, I examined the different foraging strategies for the different macronutrient sources, pollen and nectar, and the inter-individual variation in bee foraging performance. I also looked at how stressors, such as pesticides, can impact on bee foraging efficiency. I compared two social Hymenoptera that vary in their level of social complexity: the European honey bee (Apis mellifera L.) and the buffed-tailed bumblebee (Bombus terrestris L.). I used Radio Frequency Identification (RFID) to automatically track the foraging behaviour of bees throughout their life. I found that honey bee and bumblebee colonies rely on a subset of very active bees to supply the whole colony needs. In honey bees, these foragers are more efficient and collect more pollen. I also identified different strategies for pollen or nectar collection in both species. Using manipulative experiments, I then showed that bees exhibit consistent inter-individual different behaviours in a spatial learning task and that pesticides impair visual learning. My thesis aims at better explaining the causes of vulnerability of pollinators to sublethal pesticides and other environmental stressors. The results highlight the need for considering behavioural diversity as an adaptation for social insects, as well as a potential dimension of colony-level vulnerability to environmental stressors that can impair the whole colony nutritional balance.

Many bumblebee species are suffering from the effects of habitat fragmentation and population isolation. In some cases, populations have lost genetic diversity due to genetic drift and it is possible they are now at heightened risk of extinction. Inbreeding may be particularly costly to bumblebees because, as Hymenoptera, their complementary sex determination system can lead to the production of sterile or inviable diploid males. However, little is known about the effect that diploid male production has on bumblebee colony fitness. Here, the consequences of brother-sister mating in the bumblebee Bombus terrestris are investigated, and the production of diploid males was found to exert considerable costs at the colony level by reducing productivity and survival. Diploid males may therefore act as indicators of the genetic health of populations, and their detection could be used as an informative tool in hymenopteran conservation. Due to the costs associated with inbreeding, selection may have favoured the evolution of kin recognition systems in bumblebees. Data are presented that suggest that B. terrestris can discriminate between kin and non-kin as gynes were less willing to mate with siblings compared to non-relatives. Theory predicts that inbreeding may impose further costs on bumblebees through increased levels of parasitism, but empirical data are scarce. The relationship between population genetic diversity and parasite prevalence is assessed using Hebridean island populations of Bombus muscorum and Bombus jonellus. In the more outbred B. jonellus, there was no relationship between parasite prevalence and population heterozygosity. But prevalence of the gut parasite Crithidia bombi and the tracheal mite Locustacarus buchneri were found to be higher in populations of B. muscorum that had lower genetic diversity. In addition to assessing infection status, the activity of the immune system was assessed in each individual bee. However, there was no relationship between population heterozygosity and these immune parameters. This suggests that, in some Hymenopteran species, as populations lose genetic diversity the impact of parasitism will increase, potentially pushing threatened populations closer to extinction. Therefore, preventing population fragmentation by the creation of suitable habitats and by ensuring connectivity between habitat patches are important aspects of hymenopteran conservation. Finally, this thesis investigates the potential threat of pathogen spread from commercially reared bumblebees used for crop pollination to wild bumblebees. Although no direct evidence for parasite spillover is found, the prevalence of C. bombi was significantly higher in B. terrestris by the end of the season on farms that used commercial bumblebees compared to farms that did not. This high prevalence does suggest that pathogen spillover is a potential threat and it would be preferable to reduce the usage of commercial bumblebees where possible. For example, sowing wild flower mixes could boost natural pollinator populations, which in turn would benefit soft fruit pollination. Overall, this thesis contributes to our knowledge of the consequences of inbreeding in bumblebees and the relationship between genetic diversity and parasite prevalence. It provides a greater understanding of the factors that might be pushing threatened pollinators towards extinction and as a whole provides important information that may inform conservation practitioners, whose aim is to protect the future of our hymenopteran pollinators.

Bumblebees have shown both long and short-term declines throughout their range. These declines may be attributed to a range of factors including changes in land use, alterations in climatic conditions and species introductions. However, management strategies for bumblebee conservation often focus on provision of summer forage resources and other factors are frequently overlooked. Provision of spring forage and nesting sites for bumblebee queens are rarely considered, though colony foundation and early colony growth are two of the most sensitive stages in bumblebee life history. Here, the efficacy of certain agri-environment prescriptions for providing spring forage and nest sites for bumblebees is assessed, highlighting a need for specific schemes targeted towards the provision of these vital resources in the rural environment. The nesting ecology of bumblebees is poorly understood because wild colonies are difficult to locate. However, a greater knowledge of the colony-level effects of environmental change is crucial to understanding bumblebee declines. Attracting bumblebee queens to nest in artificial domiciles could provide a valuable tool for studying colony-level responses. However, domicile trials and the findings of a literature review presented here demonstrate that this approach may be largely impractical for use in the UK. Conversely, a nationwide public bumblebee nest survey produced numerous data regarding nest site preferences among bumblebee species and also demonstrated that citizen science may also provide a sensitive method for detecting declines in currently common bumblebee species. An understanding of the ecology of species interactions and coexistence can provide valuable insights into factors that may influence declines. Data presented here suggest that coexistence between some bumblebee species may be maintained by resource partitioning based on diel activity patterns that are linked to species-specific environmental tolerances. If this is the case, the potential role of climate change in bumblebee declines may be severely underestimated. There is also increasing evidence that genetic factors may play a role in bumblebee losses, accelerating declines of small, fragmented populations as a result of reduction in genetic diversity and inbreeding depression. Here, the feasibility of reintroducing British B. subterraneus (now extinct in the UK) from New Zealand into England is assessed using population genetic techniques. The findings suggest that the population history of B. subterraneus in New Zealand has resulted in a dramatic loss of genetic diversity and high genetic divergence from the original UK population, suggesting that it may not be a suitable for use in the reintroduction attempt. This work draws together some understudied aspects of bumblebee ecology with a particular focus on nest site requirements, availability of spring forage, mechanisms of avoidance of inter-specific competition and population genetic processes. The potential role of these in bumblebee declines is considered and new data relevant to the conservation of these important species is presented. It is hoped that this work will inform future management strategies for bumblebee conservation, highlight areas in need of further study and provide a sound starting point for future research in these areas.

L’érosion de la biodiversité prairiale est devenue une préoccupation majeure à l’échelle Européenne. Le pâturage ovin est supposé avoir un impact négatif sur la diversité prairiale, en raison de leur forte sélectivité pour les dicotylédones, indispensables aux insectes pollinisateurs. Comparés aux bovins, les ovins structurent peu les couverts et créent également moins de niches écologiques contrastées. L’objectif de cette thèse était de tester la faisabilité, et les bénéfices en pâturage ovin, d’une conduite en rotation dans laquelle une sous-parcelle est temporairement exclue du pâturage au moment du pic de floraison. Les effets de cette conduite ont été comparés à ceux d’un pâturage continu au même chargement. Au delà des indicateurs directs de performances zootechniques et de biodiversité (plantes, papillons, bourdons et carabes), nous avons analysé la sélection alimentaire des brebis dans les deux modes de conduite. Nous avons aussi cherché à appréhender comment la race et la fertilité du milieu pouvaient moduler la faisabilité d’un tel pâturage tournant et son intérêt vis-à-vis de la préservation de la biodiversité. Indépendamment de leur race, les brebis ont présenté une sélection alimentaire accrue vis-à-vis des dicotylédones dans les parcelles pâturées en rotation, qui a rapidement entraîné une diminution de leur richesse floristique en comparaison des parcelles pâturées en continu. L’augmentation de l’intensité de floraison des sous parcelles temporairement exclues de la rotation a favorisé les bourdons, probablement en raison de l’augmentation de la ressource en pollen et en nectar. En revanche, ce mode de gestion n’a pas permis d’augmenter la densité ni la richesse spécifique des papillons et des carabes. Le bénéfice d’une mise en défens temporaire d’une partie des parcelles semble donc moindre qu’en pâturage bovin. Définir les dates de mises en défens par rapport à la floraison d’espèces indicatrices, moduler la durée de la mise en défens en fonction de la pousse de printemps, et prolonger l’exclusion de certaines parcelles en automne et en hiver sont autant de pistes qu’il nous reste à explorer, afin de déterminer les conditions d’application optimale d’une telle conduite. / Biodiversity loss in grasslands is a major concern across Europe. Sheep grazing is rarely considered the best method for delivering conservation objectives, as the result of their strong diet selection on forbs and legumes, which in turn negatively impacts nectar-dependent insect groups. Compared with cattle, sheep also produce a strong homogeneization effect and reduce habitat diversity. The objective of this thesis was to analyze the feasibility and environmental benefits of a rotational grazing management, in which sheep were temporarily excluded from a sub-plot at flowering peak. It was compared with continuous grazing in the same stocking rate. In addition, to measurements of animal performances and biodiversity indicators (plants, butterflies, bumblebees and ground beetles), we analyzed diet selection by ewes in the two grazing managements, and how sheep breed and soil fertility could modulate biodiversity outputs in the rotational management. Independently of sheep breed, the ewes increased their selection of forbs and legumes in rotationally-grazed plots. This rapidly decreased plant species richness in rotationally compared with continuously-grazed plots. However, an increase of flowering intensity in temporarily ungrazed sub-plots benefited bumblebee density and species richness, as predicted by the ‘trophic level’ hypothesis. Rotational grazing management did not increase butterfly and ground beetle densities, which stresses that its benefits would be lesser than in cattle-grazed systems. Defining the exclusion period based on the flowering of indicator plant species, modulating its duration based on spring grass growth, and keeping some sub-plots ungrazed until the end of the grazing season are options that still need to be tested in order to define the optimal conditions for such a rotational grazing management.

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The ability to grip on petal surfaces is of crucial importance for the interactions between bees and flowers. In this thesis, I explore the biomechanics of attachment and morphological diversity of bee attachment devices, linking this to the behavioural ecology of bee interactions with flowers. Attachment devices come in two main kinds: claws or spines, and adhesive pads. Claw functioning is poorly described, particularly in terms of how their performance depends on body size, claw geometry, and surface roughness. Claw attachment performance was investigated using several insect species, each covering a large range of body masses. Weight-specific attachment forces decreased with body size, with claw sharpness seemingly playing a role. In bees there is considerable interspecific variation in tarsal claw morphology. This variation, and arolia presence/absence, was categorised for the large bee family Apidae. Cleft/bifid claws were shown to be present in the majority of the Apidae, often with differences between sexes and clades. Using Bombus terrestris, there was no evidence that cleft claws are important for pollen collection; however, I found that the inner tooth of cleft claws can act as a backup if the main tooth breaks. Although this may be one function of cleft claws, there are clearly other unresolved functions well worth further exploration. Investigations were undertaken to explore how petal surface roughness affects bee foraging behaviour. Lab-based foraging trials on B. terrestris visiting artificial flowers varying in slope, surface texture and sugar reward revealed a trade-off between the biomechanical difficulty of visiting and handling the ‘flowers’ and the quality of the reward offered. Flowers that were difficult to grip were often avoided even if they offered a higher reward. To further investigate reward preferences of bees, the effect of sucrose concentration on honey stomach offloading times was also explored. Although the majority of petals do have a rough surface, some have slippery petals. In the field, bumblebees avoided landing on slippery hollyhock petals in favour of the easy-to-grip staminal column. In contrast, honey bees, which are smaller and have larger adhesive pads, landed on both the staminal column and the petals. Slippery petals may be an adaptation to increase contact with plant reproductive structures. Grip is also important to allow the honey bee parasite Varroa destructor to climb on to their host. Attachment forces experiments found that V. destructor could support > 300 times their body mass on honey bees, giving them strong attachment even when bees attempt to remove them through grooming. A grooming-based device for treating V. destructor was tested in an apiary trial. The device was ineffective, providing valuable information for beekeepers considering using this product. In summary, this thesis improves our understanding of the biomechanics of attachment as well as identifying several important aspects of grip in bee-plant interactions.

Bumble bees (Bombus spp.) are adept pollinators of countless cultivated and wild flowering plants, but many species have experienced declines in recent decades. Though urban sprawl has been implicated as a driving force of such losses, urban green spaces hold the potential to serve as habitat islands for bumble bees. As human populations continue to grow and metropolitan areas become larger, the survival of many bumble bee species will hinge on the identification and implementation of appropriate conservation measures at regional and finer scales. North Texas is home to some the fastest-growing urban areas in the country, including Denton County, as well as at least two declining bumble bee species (B. pensylvanicus and B. fraternus). Using a combination of field , molevular DNA and GIS methods I evaluated the persistence of historic bumble bee species in Denton County, and investigated the genetic structure and connectivity of the populations in these spaces. Field sampling resulted in the discovery of both B. pensylvanicus and B. fraternus in Denton County's urban green spaces. While the relative abundance of B. fraternus in these spaces was significantly lower than historic levels gleaned from museum recors, that of B. pensylvanicus was significantly higher. Statistical analyses found that both bare ground and tree cover surrounding sample sites were negatively associated with numbers of bumble bee individuals and hives detected in these green spaces. Additionally, limited genetic structuring of bumble bee populations was detected, leading to the conclusion that extensive gene flow is occurring across populations in Denton County.

No description available.