Forest clearings can potentially be an important resource for pollinators. Land use history has previously been shown to be important for the number of species and individuals of butterflies in clearings, with a larger number where it historically was meadows, even after a generation of production forest. Plant species richness has also been shown to be influenced by land use history. In this study, pollinators were collected with pan traps on clear-cuts that had at least one generation of production forest, which in the 1870s was either meadow or forest. The results showed that there was no difference in species numbers and individual number between the two types of clear-cuts when it comes to Bombus spp, Syrphinae spp, Lepturinae spp, Cetoniidae spp and Trichius spp. The result is surprising because there is a greater frequency of herbs in clearings that previously had been meadow and the expectations was therefore that there should be more pollinators there. There are indications that the catchability when it comes to the pan trap method is inversely proportional to the floral richness, and the lack of differences could therefore be due to sampling bias.

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.

No description available.

Bumblebees have undergone dramatic declines both in Britain and further afield during the last century. Bumblebees provide a crucial pollination service to both crops and wild flowers. For these reasons, they have received a great deal of research attention over the years. However, the ecology of wild bumblebee nests and the interactions between nests and other species, particularly vertebrates has been somewhat understudied. This is largely due to the difficulty in finding sufficient nests for well replicated study and a lack of appropriate methods of observation. Here, methods for locating bumblebee nests were trialled. It was found that a specially trained bumblebee nest detection dog did not discover nests any faster than people who had received minimal instruction. Numbers of nest site searching queens provide a reliable indication of suitable nesting habitat (i.e. places where nests are more likely to be found). In order to investigate aspects of bumblebee nesting ecology wild nests were observed by filming or regular observations by either researchers or members of the public. Some buff-tailed bumblebee (Bombus terrestris) nests were collected and all the bumblebees were genotyped to identify any foreign individuals. A review of British mammalian dietary literature was conducted to identify those that predate bumblebees. Great tits (Parus major) were filmed predating bumblebees at nests and it was clear from the literature and observations that badger (Meles meles), pine martens (Martes martes) and hedgehogs (Erinaceus europaeus) predate bumblebee nests, as well as the wax moth (Aphomia sociella). No evidence for predation by any other vertebrate species was found. Behaviours recorded included parasitism by Psithyrus, apparent nectar theft and possible usurpation by true bumblebees, egg-dumping by foreign queens and drifting and drifter reproduction by foreign workers. These events may cause harm to colonies (for example, through horizontal transmission of pathogens, or exploitation of the host nest’s resources). Alternatively where for example, usurpation by true bumblebees, egg-dumping or drifting is successful, these alternative reproductive strategies may increase the effective population size by enabling a single nest to produce reproductives of more than one breeding female. These data found that wild buff-tailed bumblebees (B. terrestris) nests with a greater proportion of workers infected with Crithidia bombi were less likely to produce gynes than those with fewer infected workers. Gyne production also varied dramatically between years. There is a growing body of evidence that a class of frequently used insecticides called neonicotinoids are negatively impacting bumblebees. An experiment was conducted using commercial colonies of buff-tailed bumblebees (B. terrestris) which were fed pollen and nectar which had been treated with the neonicotinoid imidacloprid at field realistic, sub-lethal levels. Treated colonies, produced 85-90% fewer gynes than control colonies. If this trend is representative of natural nests feeding on treated crops, for example, oilseed rape and field beans or garden flowers, then this would be expected to cause dramatic population declines. In this thesis methods for locating bumblebee nests have been tested, new behaviours have been identified (for example, egg-dumping by queens and predation by great tits) and estimations for rates of fecundity and destruction by various factors have been provided. Doubt has been cast over the status of some mammals as predators of bumblebee nests and estimates for gyne production, nest longevity, etc, have been given. More work is needed, especially observations of incipient nests as this is when the greatest losses are thought to occur.

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

Bumble bees (Bombus spp.: Apidae) are valuable pollinators of many crop and wildflower species. However, in some situations their potential is limited. Evaluation of, and management to improve bumble bee efficacy should include spatial information which is currently limited. Distance and direction determine the success of gene flow via pollen cross-over within and between plant populations at several scales. Studies of movement by bumble bees at large scales in semi-natural and intensively managed habitats are scarce. Few studies of bumble bee dispersal from the nest exist, particularly in relation to crops. At a small scale, directional rather than random movement between flowers has benefits for pollen flow. Results to date of directionality studies at small scales and their interpretation are inconsistent. The purpose of this thesis was to assess distances and directions moved by foraging bumble bees at a range of scales in two contrasting farm habitats in order to predict their pollination potential. A novel method was developed to mark automatically all the occupants of nests of bumble bees B. terrestris (L.) placed around a Lucerne seed crop Medicago sativa L. in New Zealand. Reobservation data from eight nests showed that of bumble bees which foraged within the crop, 81 % travelled ≤ 50 m and 56% ≤ 20 m from their nest. Results should be interpreted with extreme caution because fewer than 1 % of bumble bees marked at nests were reobserved in the crop. Because it was not established where the other 99% of the bumble bees went, foraging areas for nests could not be calculated as anticipated. Theories to explain the non-specificity of bumble bees to the crop include; resource depletion near nests, competition with honey bees in the crop, or an evolved strategy to disperse in order to minimise nest predation. Lucerne flowers contained a significantly lower concentration of sugar in nectar, and significantly fewer pollen grains than did those of purple loosestrife Lythrum salicaria L., a species on which bumble bees appeared to forage in preference Lucerne. The higher rewards offered by L. salicaria may have diverted bumble bees from the less-rewarding Lucerne crop. In a Norwegian meadow system, all foraging bouts by bumble bees B. lucorum (L.) within a patch of wood cranesbill flowers Geranium sylvaticum L. were random with respect to direction. This result is not consistent with predictions, based on optimal foraging theory, that movement should be directional to enable optimal pollen flow, and to avoid revisitation of just-emptied flowers by the pollinator. A medium-scale study of several bumble bee species moving between patches of northern wolfsbane Aconitum septentrionale Koelle in Norway revealed considerable loyalty by bumble bees to patches in which they were marked. In a different landscape-scale study (over 5 ha), several bumble bees exhibited a high degree of loyalty to areas in which they were marked (87% were reobserved ≤ 50 m from marking points). These restricted movement patterns are discussed in terms of potential pollen flow. Of 260 bumble bees marked, only five were recorded crossing between meadows, which could be a result of innate loyalty to small forage areas, an artefact of the sampling technique used, or forest boundaries acting as physical impediments to movement. In the future, spatial data of the type collected in this thesis will aid in the management of bumble bee populations to achieve both commercial and conservation goals. Spatial data can be applied to predict the optimal placement of artificially-reared nests, predict suitable isolation distances for pure seed crops, and aid in the positioning of supplementary forage sources and nest-site refuges.

Doctorat en Sciences / info:eu-repo/semantics/nonPublished

The goals of this research project were to identify the various insects observed to visit inflorescences of Echinacea angustifolia DC, and to rank these visitors according to their importance as pollinators of E. angustifolia in Saskatchewan. Studying nectar and the nectary is essential to understanding the interaction of disc florets with pollinators. Nectar-sugar production by disc florets of E. angustifolia and E. purpurea (L. Moench) was quantified from anthesis to cessation with production per disc floret peaking in the afternoon of the staminate phase (191.7 µg) and at midday of the first day of the pistillate phase (156.6 µg), respectively. Morphology of the disc-like floral nectaries of both Echinacea species was studied, as well as the ultrastructure of the nectary of E. purpurea. Modified stomata on the nectary rim are the most likely exits for nectar, but creases in the epidermis may also participate. The nectary of E. purpurea is vascularized by phloem alone, which occurred adjacent to the epidermis. Companion cells possessed wall ingrowths, and these cells may unload arriving sugar destined for either an apoplastic or symplastic pathway. Lobed nuclei were a key feature of secretory parenchyma cells, as was a predominance of mitochondria, suggesting that energy-requiring eccrine secretion predominates in E. purpurea. E. angustifolia exhibited a generalist pollination system, with pollinating insects belonging to the orders Coleoptera, Diptera, Hymenoptera, and Lepidoptera. The pollination efficiency of visitors was determined by single insect visits to bagged, virgin inflorescences followed by quantifying pollen tubes at the bases of receptive styles and/or calculating the percentage of shrivelled styles. It was determined that bumble bees (Bombus spp.) were efficient pollinators, indicating that they would likely contribute much to the pollination of E. angustifolia. Grasshopper bee flies (Systoechus vulgaris Loew) were plentiful but individually were not efficient pollinators, but taken together, they provided much pollination. Golden blister beetles (Epicauta ferruginea Say) were efficient pollinators but where yellow-petalled flowers occurred, their numbers on E. angustifolia decreased. Honey bees (Apis mellifera L.) were efficient pollinators and were present in low numbers without managed introduction. Pierid (2003) butterflies were regular visitors and efficient pollinators, and likely contributed significantly to E. angustifolia pollination. When introduced, the alfalfa leafcutter bee (Megachile rotundata Fabr.) preferred not to forage on E. angustifolia and as such, these solitary bees were not suitable as managed pollinators. In large agricultural plantings of E. angustifolia, however, native insects may not be capable of providing sufficient pollination for seed production when floral competition occurs.

The goals of this research project were to identify the various insects observed to visit inflorescences of Echinacea angustifolia DC, and to rank these visitors according to their importance as pollinators of E. angustifolia in Saskatchewan. Studying nectar and the nectary is essential to understanding the interaction of disc florets with pollinators. Nectar-sugar production by disc florets of E. angustifolia and E. purpurea (L. Moench) was quantified from anthesis to cessation with production per disc floret peaking in the afternoon of the staminate phase (191.7 µg) and at midday of the first day of the pistillate phase (156.6 µg), respectively. Morphology of the disc-like floral nectaries of both Echinacea species was studied, as well as the ultrastructure of the nectary of E. purpurea. Modified stomata on the nectary rim are the most likely exits for nectar, but creases in the epidermis may also participate. The nectary of E. purpurea is vascularized by phloem alone, which occurred adjacent to the epidermis. Companion cells possessed wall ingrowths, and these cells may unload arriving sugar destined for either an apoplastic or symplastic pathway. Lobed nuclei were a key feature of secretory parenchyma cells, as was a predominance of mitochondria, suggesting that energy-requiring eccrine secretion predominates in E. purpurea. E. angustifolia exhibited a generalist pollination system, with pollinating insects belonging to the orders Coleoptera, Diptera, Hymenoptera, and Lepidoptera. The pollination efficiency of visitors was determined by single insect visits to bagged, virgin inflorescences followed by quantifying pollen tubes at the bases of receptive styles and/or calculating the percentage of shrivelled styles. It was determined that bumble bees (Bombus spp.) were efficient pollinators, indicating that they would likely contribute much to the pollination of E. angustifolia. Grasshopper bee flies (Systoechus vulgaris Loew) were plentiful but individually were not efficient pollinators, but taken together, they provided much pollination. Golden blister beetles (Epicauta ferruginea Say) were efficient pollinators but where yellow-petalled flowers occurred, their numbers on E. angustifolia decreased. Honey bees (Apis mellifera L.) were efficient pollinators and were present in low numbers without managed introduction. Pierid (2003) butterflies were regular visitors and efficient pollinators, and likely contributed significantly to E. angustifolia pollination. When introduced, the alfalfa leafcutter bee (Megachile rotundata Fabr.) preferred not to forage on E. angustifolia and as such, these solitary bees were not suitable as managed pollinators. In large agricultural plantings of E. angustifolia, however, native insects may not be capable of providing sufficient pollination for seed production when floral competition occurs.

Plants often occur aggregated into clusters and this spatial pattern is supposed to affect pollinator behaviour and pollen dispersal. Such pollinator reaction may influence reproductive success of zoogamous plant species both in terms of number of available mates and their genetic diversity (nearby growing plant individuals are also often closely related, especially in clonal plants). In the present thesis, I investigated the influence of plant spatial aggregation on pollinator behaviour and how this translates into pollen transfer. For that purpose, I carried three experiments. In the Experiment 1, I used potted plants placed into arrays and aggregated into four patches in order to track the pollen dispersal by means of a UV-dye pollen analogue. I manipulated distances between plants within clusters (dense × loose) and between clusters (near × far). I conducted this experiment for three plant species differing in their pollinator spectra. In the Experiment 2, I observed pollinator foraging sequences (sequences of visited plant individuals) under the same experimental design as for the first experiment, but I carried out this experiment for five plant species. In addition in one study species, Dianthus carthusianorum, I conducted the Experiment 3 to get better insight into pollination effectiveness...