Summary1. Generalist herbivorous insects, which feed on plant tissue that is nutritionally heterogeneous or varies in its content of secondary metabolites, often benefit from dietary mixing through more balanced nutrient intake or reduced exposure to harmful secondary metabolites. Pollen is similarly heterogeneous as other plant tissue in its content of primary and secondary metabolites, suggesting that providing their offspring with mixed pollen diets might be a promising strategy for pollen generalist bees to complement nutrient imbalances or to mitigate harmful secondary metabolites of unfavourable pollen. 2. In the present study, we compared larval performance of the pollen generalist solitary bee species Osmia cornuta (Megachilidae) on five experimental pollen diets that consisted of different proportions of unfavourable pollen diet of Ranunculus acris (Ranunculaceae) and favourable pollen diet of Sinapis arvensis (Brassicaceae). In addition, we microscopically analysed the pollen contained in the scopal brushes of field-collected females of O. cornuta and three closely related species to elucidate to what degree these pollen generalist bees mix pollen of different hosts in their brood cells. 3. In striking contrast to a pure Ranunculus pollen diet, which had a lethal effect on most developing larvae of O. cornuta, larval survival, larval development time and adult body mass of both males and females remained nearly unaffected by the admixture of up to 50% of Ranunculus pollen diet to the larval food. 4. Between 42% and 66% of all female scopal pollen loads analysed contained mixtures of pollen from two to six plant families, indicating that pollen mixing is a common behaviour in O. cornuta and the three related bee species. 5. The present study provides the first evidence that the larvae of pollen generalist bees can benefit from the nutrient content of unfavourable pollen without being negatively affected by its unfavourable chemical properties if such pollen is mixed with favourable pollen. We conclude that the widespread pollen mixing by females of pollen generalist bees should also be considered as a possible strategy to exploit flowers with unfavourable pollen and to optimize larval food quality.
Within the genus Osmia, the three subgenera Osmia, Monosmia, and Orientosmia form a closely-related group of predominantly pollen generalist ('polylectic') mason bees. Despite the great scientific and economic interest in several species of this clade, which are promoted commercially for orchard pollination, their phylogenetic relationships remain poorly understood. We inferred the phylogeny of 21 Osmia species belonging to this clade by applying Bayesian and maximum likelihood methods based on five genes and morphology. Because our results revealed paraphyly of the largest subgenus (Osmia s.s.), we synonymized Monosmia and Orientosmia under Osmia s.s. Microscopical analysis of female pollen loads revealed that five species are specialized ('oligolectic') on Fabaceae or Boraginaceae, whereas the remaining species are polylectic, harvesting pollen from up to 19 plant families. Polylecty appears to be the ancestral state, with oligolectic lineages having evolved twice independently. Among the polylectic species, several intriguing patterns of host plant use were found, suggesting that host plant choice of these bees is constrained to different degrees and governed by flower morphology, pollen chemistry or nectar availability, thus supporting previous findings on predominantly oligolectic clades of bees.
Summary1. Pollen host choice in bees is in many cases highly conserved, which might partly be due to physiological limitations of bee larvae to digest non-host pollen. These limitations need to be overcome in order to incorporate new pollen hosts; however, the mechanisms underlying such host expansion are poorly understood. 2. In this study, we examined intra-and interpopulational variation in the ability of larvae of the solitary bee species Osmia cornuta (Megachilidae) to develop on a non-host pollen diet of Ranunculus acris (Ranunculaceae) by comparing larval performance within and between five geographically distant European populations. 3. The majority of bee larvae from all tested populations died when reared on the Ranunculus pollen diet. Between 10% and 43Á5% of all larvae per population reached the cocoon stage, and 48% of these emerged as viable adults from the cocoons, indicating that the physiological ability to cope with the unfavourable properties of Ranunculus pollen exists in each population. 4. The bee larvae of one population exhibited significantly reduced survival on the Ranunculus pollen diet compared with three of the four other populations. 5. Although bees that successfully developed on the Ranunculus pollen diet showed a distinctly prolonged development time, exhibited higher mortality during diapause and reached a considerably lower adult weight compared with individuals fed the control pollen diet, several of the Ranunculus fed individuals were able to reproduce and to sire viable offspring. 6. This study provides the first evidence for both intra-and interpopulational variation in the physiological ability of solitary bees to digest non-host pollen. This variation might enable host expansion and subsequent host shifts in response to natural selection.
Empirical evidence suggests that pollen chemistry plays an important role in shaping the pollen host spectra of many bee species. Although the underlying mechanisms are poorly understood, pollen diets of several plant taxa have experimentally been found to impede larval development of unspecialized bees. The pollen of all plant taxa, for which such a detrimental effect on bee larval development has been observed so far, is freely accessible in the flowers and thus easily harvestable for flower visitors, suggesting that this pollen might be chemically protected in order to reduce its loss to pollen-feeding animals. In the present study, we compared larval performance of five solitary bee species on pollen diets of the two Fabaceae species Onobrychis viciifolia and Lotus corniculatus, which have their anthers concealed inside the flowers, with that on control diets composed of host pollen provisions. As the complex flower morphology of the two Fabaceae species already considerably narrows the spectrum of pollen harvesting bee taxa, which might supersede costly chemical protection of the pollen, we expected bees that usually do not exploit Fabaceae to develop well on Onobrychis and Lotus pollen diets. Larval survival on the Onobrychis pollen diet was successful for all five bee species tested. In contrast, larval survival on the Lotus pollen diet was reduced in three species despite the fact that Lotus flowers are more difficult to exploit for pollen than Onobrychis flowers. We conclude that there is no trade-off between pollen concealment and pollen defence in Lotus and that pollen of morphologically complex flowers with a restricted visitor spectrum is not necessarily an easy-to-use nutritional source.
The availability of nesting resources influences the persistence and survival of bee communities. Although a positive effect of artificial nesting structures has frequently been shown for aboveground cavity-nesting wild bees, studies on below ground-nesting bees are rare. Artificial nesting hills designed to provide nesting habitats for ground-nesting bees were therefore established within the BienABest project in 20 regions across Germany. Wild bee communities were monitored for two consecutive years, accompanied by recordings of landscape and abiotic nest site variables. Bee activity and species richness increased from the first to the second year after establishment; this was particularly pronounced in landscapes with a low cover of semi-natural habitat. The nesting hills were successively colonized, indicating that they should exist for many years, thereby promoting a species-rich bee community. We recommend the construction of nesting hills on sun-exposed sites with a high thermal gain of the substrate because the bees prefer south-facing sites with high soil temperatures. Although the soil composition of the nesting hills plays a minor role, we suggest using local soil to match the needs of the local bee community. We conclude that artificial nesting structures for ground-nesting bees act as a valuable nesting resource for various bee species, particularly in highly degraded landscapes. We offer a construction and maintenance guide for the successful establishment of nesting hills for bee conservation.
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