Bumble bees provide vital pollination services in both native and agricultural landscapes. However, in recent years, bumble bee populations have experienced global population declines. The primary causes of these declines have been attributed to the environmental impacts of pathogens, pesticide use and habitat fragmentation. While research has examined the impacts of pathogens, there is limited information on the effects of pesticides and habitat fragmentation on native bumble bees. Hence, the objectives of my dissertation research were to: 1) assess the toxicological impacts of pesticides used in two important bee-pollinated crops on queens and workers; 2) determine the impacts of forage resource availability on bumble bee colonies; 3) examine pollen foraging behavior of bumble bees in a late season mass-flowering agricultural landscape; and 4) document observations on trends towards bivoltinism in three western North American bumble bees. This research was conducted in the lab using wild and lab reared colonies, and in an agricultural landscape in the Willamette Valley of Western Oregon. In pesticide bioassays the impacts of residual toxicity of five classes of pesticides used in highbush blueberry and red clover cropping systems, were tested on queen and worker bumble bees, respectively. The results indicated variation in responses to the same insecticide by queens and workers, and to the same class of compounds by workers. Also, toxic effects were documented for both queens and workers to pesticides considered to be "safe" for bees. The impacts of forage resource availability were evaluated by exposing bumble bee colonies to four quantities of pollen, four quantities of nectar and three feeding frequencies of pollen. The study documented an inverse relationship between larvae and workers to resource type and availability. In addition, the type of larval mortality displayed by colonies, larval ejection or within clump mortality, was dependent on worker mortality. The individual and colony-level pollen foraging behavior of bumble bees was examined by placing colonies in red clover. Observations in the field on forager abundance, and at the colony-level on the duration and number of pollen trips and weight of stored pollen documented that red clover is an important resource for bumble bees. Pollen analysis revealed that in addition to red clover, Himalayan blackberry was also a key forage resource for bumble bees. Red clover resources at the end of the season may also benefit bumble bees by allowing for the creation of a second generation. Observations on both field and lab-reared queens document a trend towards bivoltinism in three species of western North American bumble bees. Agricultural habitats are vital for sustaining bumble bee populations. However, given the potential for pesticide impacts and temporal availability of flowering plants, these landscapes must be managed to provide maximum benefit to bumble bees. Results from this research should assist growers and researchers in developing landscape management and production practices geared toward the conservation and enhancement of native Bombus spp. populations in western Oregon. / Graduation date: 2011

No description available.

The distribution and abundance of six bumblebee species (Bombus) and their associated flowers were studies for 12 weeks in 26 discrete subalpine meadows over the summer of 1981. Principal component analysis of flower densities identified phenological patterns of blooming and differential affinities of plants for moisture as the major contributors to flower variation among the meadows. A positive relationship between bumblebee species diversity and meadow area in the early summer and mark-recapture experiments suggested that queens are attracted to large meadows for nest establishment. After colony initiation meadow floristics govern Bombus species diversity and abundance. Results showed that: (1) mid- to late-summer bumblebee diversity, the temporal occurrence of new queens, and the densities of two species were correlated with the density of nectar producing flowers; (2) flower composition in meadows where local extinctions of at least one Bombus species occurred differed from those where all species persisted; and (3) the number of species reaching reproductive fruition was independent of bumblebee species composition. These results underscore the importance of nectar resources in Bombus colony and guild development. Multiple regression analyses were used to relate the effects of meadow size, elevation, and floristics to observed densities of Bombus flavifrons and B. rufocinctus. In mid- to late summer a significant portion of the variation in densities not explained by these variables was accounted for by densities of congenerics. In meadows where all Bombus were removed except B. flavifrons or B. rufocinctus significant positive density compensation was observed for the latter, but not the former. These results are suggestive of relationships among bumblebee densities, intensity of competition, colony ontogeny, colony reproduction, and local floristics that question both the (1) simplistic views of Bombus guild organization based on the paradigm of tongue length, corolla depth, and character displacement; and (2) the stochastic view that flowering phenologies and developmental time lags thwart resource-based guild organization.

Locomotion is central to the survival of many animal species; however large variation in performance, for example in speed or endurance, exists between individuals within a species. Using the bumblebee species, Bombus impatiens, I studied the extent of the variation in several flight performance traits and how they are associated. I first addressed how bumblebee workers vary in foraging effort and observed that only around half of the monitored individuals underwent foraging activity. Additionally, significant variation in metabolic rate between foragers and non-foragers was uncovered. I further investigated if such variation could be associated with flight performance capacity, such as an individual’s ability to carry a load, their flight speed and distance traveled, their wing morphology and kinematics, and their flight metabolic rate. These traits are commonly measured to characterize flight capacity in insects, however the links between them have yet to be investigated. Links between morphology, wing kinematics and peak metabolic rate previously uncovered in the literature were observed in my analysis, although variation in their scaling with body mass was detected. Vertical force scaled isometrically with body mass but was not related to it when expressed in on a mass specific basis (VF m-1g-1, where m is gravitation acceleration). In regard to forward flight speed, body mass does have an affect, however it alone does not have a great degree of explanatory power and other factors such as morphology and wing kinematics are likely to play a greater part in its determination. Finally, maximum flight speed had a significant relationship with total flight time. Together, these results demonstrate that some links do exist between flight performance traits, however links are not present between all traits and certain flight performance traits should be treated as independent of each other.

The development of three colonies of Bombus fervidus (Fabricius) and two of Bombus pennsylvanicus (Degeer) was studied. Population levels and the production of males and young queens varied among colonies and between years. Mean longevity of worker bees decreased significantly toward the end of the season. Mean size of emerging worker bees increased significantly with time. The decreased survivorship may be related to the seasonal size increase of the workers, which would result in an increased proportion of foragers. Age-specific life tables and survivorship curves were developed for workers within colonies, and for workers in cohorts based on emergence date. An increase in brood developmental times occurred near the end of the colony cycle. In the two B. pennsylvanicus colonies, oviposition of fertilized eggs decreased during the transition period from worker to queen production, and large numbers of male-producing eggs were laid. At the time of queen production a sudden increase in size of emerging females occurred. Neither a gradual nor a sudden change in larva/worker ratio appears sufficient to explain caste determination. Rates of parasitization of bumble bee queens at three sites in Southwestern Virginia were studied. Endoparasites of spring queens included Locustacarus buchneri (Stammer) (Acarina: Podapolipidae), a mite infesting the abdominal air sacs; the nematode Sphaerularia bombi Dufour (Tylenchida: Nematoda), and gregarious braconid larvae, probably Syntretus sp. (Hymenoptera: Braconidae). Ectoparasites included hypopi of the mite Kuzinia americana Delfinado and Baker (Acari: Acaridae) and Parasitus spp. mites (Acari: Parasitidae). / M.S.

Over 80% of wild angiosperms are reliant upon animal pollination for fruit and seed set and bees and other insects provide a vital pollination service to around a third of the crops we produce. Habitat loss, climate change and disease spread all threaten pollinator populations, with local declines and range contractions in honeybees and bumblebees leading to concerns that crop production may suffer as a result of pollinator shortages. Whilst agriculture and wildlife are often presented as being at odds with one another, the relationship between farmers growing pollination dependent crops, and the bees and insects that service them could be mutualistic. Flowering crops planted by farmers can provide an important source of forage to wild bees, whilst in return wild bees can contribute to ensuring farmers achieve adequate yields of marketable crops. The potential of this mutualistic relationship can be maximised by farmers by adopting management practices that reduce harm to, and enhance the wellbeing of, the wild bees around their farm. A group of common pesticides (neonicotinoids) used by farmers have recently been linked to pollinator ill health. Sub-lethal effects resulting from exposure to the neonicotinoid imidacloprid have been reported in honeybees and bumblebees, with bumblebee reproductive success found to diminish as a result of exposure to field realistic doses of this agrochemical. Here, the mechanism behind the reduced queen production in bumblebee colonies is suggested, with bees exposed to imidacloprid showing reduced efficiency in foraging for pollen. Farmers dependent upon pollinating insects for crop production can opt to avoid the use of pesticides known to harm these insects, however future studies are needed to identify safer alternatives that can be use in their place. Farmers can choose to increase the number of bees at their farms by utilising domesticated honeybees and purchasing commercially reared bumblebees. The use of these pollinators can ensure a minimum number of bees in the vicinity of a crop, and facilitate the production of crops at times when wild bee numbers are low. Concerns have been raised, however, regarding the use of commercially reared bees, mostly in regard to pathogen and parasite transmission, but also in respect to the possibility of outcompeting native species. Here the frequency and severity of attacks on commercial Bombus terrestris colonies, by the wax moth, an understudied bumblebee pest, are examined. Wax moths were found to infest almost half of the bumblebee nests deployed at fruit farms, with around a third of infestations resulting in nest destruction. Farmers investing in commercial bees will want to reduce the impact of harmful pests that may result in a reduced pollination service being delivered. Wax moth infestation rates at the study farms using commercial bees were high and the potential of a ‘spill- back’ effect on wild bees was examined. No evidence was found to suggest that nests in close proximity to these farms were any more or less likely to suffer from an attack than nests situated further away. Nest size was found to be the most significant predictor of an infestation, with larger nests more prone to wax moth attacks. Whilst farmers can utilise domesticated and commercially reared bees, relying on one source of pollination is inherently risky, and the most robust service will likely be provided by a range of pollinators. As well as reducing the use of chemicals known to harm beneficial insects, farmers can improve the habitat around their farms to help encourage and sustain wild pollinator populations. Sowing flower strips has been found to increase the abundance and diversity of pollinating insects, however, studies linking the use of these strips to crop production are lacking. Here we demonstrate for the first time that sowing small flower strips, adjacent to strawberry crops serviced by both wild and managed bees, can increase the overall number of pollinators foraging on the crop. This thesis contributes to our understanding of the implications of farm management decisions on pollinator health. It provides experimentally based evidence to guide farmers in making informed decisions regarding the future of crop pollination services and highlights the need for an integrated approach to managing pollination services for sustainability.

Bumblebees are ecologically and economically important as pollinators, but some species are suffering severe declines and range contractions. In this thesis, three cryptic bumblebee species are studied to elucidate differences in their distribution, ecology and population genetics. As a result of their high morphological similarity, very little is known about the lucorum complex species: B. lucorum, B, cryptarum and B. magnus. In this study, their distributions across Great Britain were assessed using molecular methods, revealing that B. lucorum was the most abundant and most generalist of the three species, whereas B. magnus was the rarest and most specialised, occurring almost exclusively on heathland. Additionally, both B. magnus and B. cryptarum were more likely to be present at sites with cooler summer temperatures. Cryptic species represent interesting models to investigate the levels of niche differentiation required to avoid competitive exclusion. Characterising the niches of these species at a single site across the flight season revealed differences along three niche dimensions: temporal activity, weather sensitivity and forage-resource use. These species exhibited asymmetric niche overlap; a combination of ecological divergence and spatio-temporal heterogeneity may contribute to maintaining them in sympatry. Population genetic studies can be highly informative for understanding species ecology and for conservation management. The differences in habitat specialisation exhibited by these bumblebee species provide the opportunity to test conflicting hypotheses about links between dispersal and ecological specialisation: are habitat specialists selected to have low or high dispersal ability? Based on microsatellite analysis, the generalist B. lucorum had high levels of genetic diversity and little population structure across large spatial scales. The habitat specialist B. magnus had the lowest genetic diversity but similar levels of population differentiation to the moderate generalist, B. cryptarum. However, unlike B. cryptarum, B. magnus population differentiation was not affected by geographic distance, suggesting that this specialist species may maintain effective dispersal across large scales despite being restricted to a fragmented habitat. Bergmann’s rule is a well-known ecogeographic rule describing geographical patterns of body size variation, whereby larger endothermic species are found more commonly at higher latitudes. Ectotherms, including insects, have been suggested to follow converse Bergmann’s gradients, but the facultatively endothermic nature of bumblebees makes it unclear which pattern they should adhere to. This thesis reports caste-specific differences in body size between the three lucorum complex species in agreement with Bergmann’s rule: queens and males of B. cryptarum and B. magnus, which were found more commonly at higher latitudes and at sites with cooler temperatures, were larger than those of B. lucorum. Population genetic studies of invertebrates generally require the destruction of large numbers of individuals, which is often undesirable. Testing a variety of faecal collection and DNA extraction methods demonstrated that it is possible to obtain DNA of sufficient quality for genotyping from bumblebee faeces, without harming the individuals. This method would be valuable for studies of rare or declining bee species, for queens in reintroduction projects, and may be applicable to other arthropods. Overall this thesis contributes substantially to our knowledge of the ecology and population genetics of three important pollinator species. It provides data to inform species conservation, as well as understanding of ecosystem functioning and population dynamics. Furthermore, it successfully uses these cryptic species as a model to test several fundamental ecological theories.

The amount of semi-natural grasslands has decreased in the agricultural landscape and because of this many grassland species are declining. Semi-natural grasslands are dependent on management, but different studies suggest that different management methods are most favorable to species richness and abundance. The aim of this study was to compare the effects of mowing and grazing on butterfly and bumblebee abundance in Swedish semi-natural grasslands. We used data collected through the Swedish environmental monitoring program NILS. We identified 31 comparable meadows and chose 1-10 pastures in the area surrounding each meadow. For six different species groups we calculated the number of individuals found per 100 m. By standardizing the values of the difference between the means for meadows and pastures we were able to compare the effects of the two methods on abundance in each species group. No species group showed a preference for pastures. A significant preference for meadows was found for two butterfly species groups. The four other groups showed a trend to prefer meadows before pastures. According to our study the recommended management method should be mowing, considering protection of butterflies and bumblebees.

Bees and wasps perform learning flights when departing their nest for the first few times or a newly discovered food source. Several studies have described the occurances and structure of these flights in several species, but few have examined how the insects systematically vary the characteristics of their learning flights in various conditions in order to aid the acquisition of visual information. This is best done in a species where individuals and nests can be easily manipulated and tested repeatedly. The aim of this thesis was therefore to investigate learning flights in bumblebees, where we have a good understanding of the structure and variability of flights from previous work and can design controlled experiments. I explored the similarities and differences of learning flights of workers and male bumblebees, observing their departures from the nest or an artificial flower. A second objective was to examine how differences in the learning flights affect the bumblebees’ ability to return the learnt location. The experiments were conducted inside a large greenhouse, under natural light regimes, with two large tables placed far apart, one for simulating the ground from which bees emerged when departing their nest, and the other representing a feeding site with an artificial flower. Female bumblebees performed shorter learning flights when leaving a flower than when leaving their nest, although both locations displayed similar visual scenes. At both locations, the duration and trajectory length of learning flights decreased over successive visits, but the decrease was faster at the flower location than at the nest. Bumblebees fixated both their nest and the flower during their learning flights as well as the landmarks available around the two locations, which suggests that they learned the position of the goal relative to these landmarks. When the nest and the flower were hidden and only three cylinders were shown as landmarks in tests, bees searched as accurately for the nest as for the flower. However, they were more persistent when searching for the nest than for the flower, which was not predicted from the variation of learning flights at the nest and flower locations. Another situation in which bumblebees varied the characteristics of their learning flights, but without an impact on their performance when recalling the learnt information, was after visiting flowers filled with low and high sucrose rewards. The bees performed longer learning flights after drinking at a highly rewarded flower. When departing a poorly rewarded flower, bumblebees did not fixate the flower during their learning flights. Nevertheless, the bees were able to return to both the poorly rewarded flower and the highly rewarded flower equally fast. Given the above findings, it is not evident how different durations or trajectory lengths of bumblebee learning flights might be linked to variations in learning of goal locations. Finally, I show that bumblebees of either sex decide to perform learning flights at locations that are of importance to them. Whilst the female workers always performed learning flights when departing their natal nest, the males did not and simply flew away in a straight line. However, when leaving a flower, the males did perform learning flights with characteristics similar to those of the females’ learning flights. They were also able to return to the flower, showing similar approach trajectories as workers. The thesis discusses these findings in the light of ideas and hypotheses that are linked to differential investment in learning which were observed in the various conditions here. It is also discussed why bumblebees used fixations in different ways when learning about the visual environment surrounding goals that are important to them. Whilst many results are parsimonious with the requirements for learning and active flight control to aid the acquisition of visual information, motivation also seems to play a role in varying the occurances and features of learning flights, such as seen in the bees’ greater persistence to search for their nest than for a flower.

Over the last half century, the widespread decline of bumblebees across the agricultural landscapes of Western Europe and North America has been well documented. This decline has undoubtedly been driven to a large extent by the intensification of agriculture, which has fragmented landscapes and removed large areas of suitable foraging habitat, nesting and hibernation sites. Consequently, some of the rarest Bombus species now persist only in isolated pockets of semi-natural habitat, which have been subjected to little agricultural intensification. Of the 25 Bombus species native to the UK, three species have gone extinct in recent decades and several others are severely threatened. Remaining populations of the UK’s rarest bumblebee, Bombus distinguendus, have become strongly associated with florally-rich machair grassland habitats found only in the North and West of Scotland and Western Ireland. Machair, a unique habitat that forms on soils rich in shell sand, has been maintained by rotational agricultural practices implemented by crofters. However, recent changes in crofting practices, which include the intensive grazing of machair in some areas, or conversely the abandonment of machair management all together in others, have resulted in sections of machair that have become degraded and consequently exhibit low floral abundance and species diversity. This has significant implications for species such as B. distinguendus, which have for the most part come to rely of the florally-rich swards of machair grassland. This thesis aimed to develop a greater understanding of how machair grassland habitats are utilised by foraging bumblebees, including B. distinguendus, and in turn examined the potential for restoring degraded areas of machair via a variety of methods. The research presented here examines the influence of current crofting practices on the abundance of bumblebees and their forage plant species and combines this information with a detailed exploration of the machair seed bank and potential machair restoration treatments. The specific foraging requirements of B. distinguendus were found to be similar to those of other long-tongued bumblebee species and the provision of plants from the Fabaceae family was found to be of particular importance. Current crofting practices implemented in the North and West of Scotland were, on the whole, found support low numbers of foraging bumblebees. Similarly, existing habitat management schemes, designed to provide early cover for corncrakes and foraging resources for bumblebees, were found to be largely ineffective in attracting B. distinguendus, when compared with florally-rich machair habitat. In addition, this research suggests that the existing machair seed bank is unlikely to provide a sufficient resource for reinstating florally-rich habitat to degraded areas of machair. However, this thesis has demonstrated that it is possible to implement seed mixes on machair which can reinstate species typical of machair plant communities and which also attract high numbers of foraging bumblebees. The findings of these habitat assessments and restoration trials are examined in full in the following chapters and implications for the future management of wildflower-rich machair are discussed throughout.