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Loss of natural and semi-natural habitat due to increasing human land use for agriculture and housing has led to widespread declines in bee pollinator diversity and abundance, which raised global concerns about the stability of pollination services. Bee population dynamics depend on floral resource diversity and availability in the surrounding landscape, and loss of plant biodiversity may thus directly impair the fitness of individual bee species. However, whether and how plant and resource diversity and availability affect foraging patterns, resource intake, resource quantity and nutrient quality and ultimately fitness of generalist social bees remains unclear. In this thesis, we placed hives of the Australian eusocial stingless bee Tetragonula carbonaria (Apidae, Meliponini) in natural habitat (subtropical forests) and two landscapes differently altered by humans (suburban gardens and macadamia plantations), varying in plant species richness, resource abundance and respective habitat patch size. Foraging patterns and resource intake were compared between landscapes in different seasons and colony growth and fitness were monitored over two and a half years. Bee foraging activity, pollen and sugar intake, diversity of collected pollen and resin resources, resource quantity (colony food stores), colony fitness (brood volume, queenand worker reproduction) and colony growth overwhelmingly increased with plant species richness in the surrounding habitat. However, plant species richness and thus bee fitness was highest in gardens, not in natural forests, as bees in gardens benefited from the continuous floral resource availability of both natural and exotic plants across seasons. In contrast, foraging rates and success, forager orientation and consequently colony fitness was largely reduced in plantations. While bees maximized diversity of collected resources, collecting more diverse resources did however not increase resource functionality and nutritional quality, which appeared to be primarily driven by the surrounding plant community in our study. Conversely, individual worker fitness (body fat and size) was not affected by available resource diversity and abundance, showing that colonies seem not to increase the nutritional investment in single workers, but in overall worker population size. This thesis consequently revealed the outstanding role of plant biodiversity as a key driver of (social) bee fitness by providing more foraging resources, even when only small but florally diverse patches are available.
The worldwide decline of plant and insect species during the last decades has far-reaching consequences for the functionality of ecosystems and their inherent processes. Pollination as one of them is an indispensable ecosystem service for human wellbeing. More than 85% of the worldwide flowering-plant species depend to some degree on pollination by insects (pollinators). Similarly, many pollinators depend on the flowers of the plants, as they need nectar and pollen as food resources for themselves and their offspring. However, an increasing number of pollinator and plant species are threatened by multiple, interacting, and sometimes synergistic causes (habitat loss, fragmentation, diseases, parasites, pesticides, monocultures) that are becoming a growing threat to ecosystem functioning. Given the loss of plant species diversity, it is increasingly difficult for pollinators to find food throughout the year. Therefore, this study analyses the influence of plant diversity on pollinators. The study was conducted in the course of the Jena Experiment, which is a long-term biodiversity experiment (since 2002) with 60 plant species, common to Central European Arrhenatherum grasslands. With a plant diversity gradient of 1, 2, 4, 8, 16, and 60 plant species per plot, time-series data resulted from a wide range of ecosystem processes, ranging from productivity, decomposition, C-storage, and N-storage to herbivory, and pollination. These were studied to investigate the mechanisms underlying the relationships between biodiversity and ecosystem processes.
Chapter 2 studies the spatio-temporal distribution of pollinators on flowers along an experimental plant diversity gradient. For this purpose, the pollinators were divided into four different functional groups, i.e. honeybees, bumblebees, solitary bees and hoverflies. In particular, the spatial pollinator behaviour was examined, that is, in which flowering height the flowers were visited within the plant community. In order to study the temporal component, pollinator visits were observed over the course of the day and the season. As a result, an unprecedented high resolution of plant-pollinator interactions was found. For the first time it was possible to demonstrate that the different pollinator functional groups can complementarily use different spatio-temporal niches which was most pronounced in species-rich plant mixtures,. This leads to the conclusion that species-rich plant mixtures provide sufficient resources that can be used by generalists, such as honeybees and bumblebees, as well as other pollinator functional groups, such as hoverflies and solitary bees.
Chapters 3 and 4 continues on the chemical composition of flower nectar (nectar) of various plant species. Nectar is used as food resource for adult pollinators, but is also largely used as a supply for their offspring, making it the most important pollinator reward. The chemical composition of the nectar was analysed for the two most important macronutrients, carbohydrates (C) and amino acids (AA), using high performance liquid chromatography (HPLC). Subsequently, their contents were analysed in terms of concentration, proportional content and the ratio of carbohydrates to amino acids (C:AA).
In Chapter 3, the nectar of 34 plant species from the grasslands of the Jena Experiment was compared. In doing so, similarities and/or differences of the nectar compositions were investigated with respect to the most important macronutrients carbohydrates and amino acids between the individual species but also between the most representative plant families. This should lead to a better understanding about how plant diversity influences consuming pollinators and which factors, e.g. phylogenetics, morphology or ecology, can lead to different nectar compositions. We could show that each plant species differs in terms of carbohydrate content, amino acid content and C:AA-ratio. In addition, there were clear differences between the four representative plant families Apiaceae, Asteraceae, Fabaceae and Lamiaceae regarding the proportions of essential amino acids. The proportions of the individual sugars and the C:AA-ratios also differed greatly between the four plant families. Therefore, it can be assumed that these nectar contents are family-specific. The need for differences in carbohydrate content are probably due to the different morphology of the flowers, as plants with open flowers and exposed nectar, as in Apiaceae and Asteraceae, can protect their nectar from evaporation if the nectar has a higher osmolality, which can be achieved by a higher hexose (fructose and glucose) content. Thus, the nectar can remain dilute for a longer time and consequently remain consumable for pollinators, which in turn can contribute to the pollination of plants. Fabaceae and Lamiaceae showed different results. Here the nectar was probably protected from evaporation by closed flowers, which explains the high proportion of sucrose, leading to a lower osmolality that would enhance evaporation for exposed nectar. The metabolic pathways controlling the family-specific C:AA-ratios are yet to be explored. In conclusion, it can be suggested that this study contributes to elucidating the morphological and phylogenetic characteristics that control each plant species’ nectar composition.
In Chapter 4, nectar was investigated in the context of diversity effects on the example of the plant species Field Scabious, Knautia arvensis. It was analysed to what extent the nectar quality (nutrient content) differs between plant individuals of one species. The underlying factors causing these differences in nectar composition have never been studied before. In order to investigate these coherences, plant communities in the Jena Experiment of different plant species richness levels containing the target plant species K. arvensis were used. In particular, we examined whether the nectar of K. arvensis is influenced by other neighbouring plant species, e.g. through competition for pollinators. The carbohydrate and amino acid content in nectar varied both between individuals of K. arvensis and between the different plant species richness levels. However, there were significant non-linear differences in the proportions of certain essential and phagostimulatory amino acids, which were produced proportionally more in the nectar of K. arvensis plants in species-rich plant communities, while histidine, one of the generally inhibiting amino acids tended to be less present. Our findings therefore suggest that the nectar of K. arvensis is more palatable when the plants grow in species-rich plant communities.
Overall, these studies indicate how fragile plant-pollinator interactions are but also how important plant species-rich grasslands are to support plant-pollinator interactions. Increased plant species diversity is essential to ensure the availability of flowering resources throughout the year. Pollinators, such as honeybees, bumblebees, solitary bees, and hoverflies can use the niches in time and in vertical space complementarily. However, in plant species-poor grasslands there may be more niche overlaps, which is probably due to a reduced availability of resources. This points to the need to include different plant species belonging to different plant families, whose nectar may have evolved in response to morphological flower traits and metabolic pathways. Therefore plant species diversity can supply pollinators with nectar differing in carbohydrate and amino acid content and thus differing in quality. Also C-AA ratios have proven to be a useful measurement to reveal differences between plant species. In addition, C:AA ratios were not differing in nectar of K. arvensis individuals growing in different plant species richness levels, although their nectar seemed to be more attractive in mixtures with 16 plant species, likely due to higher content of essential and phagostimulatory amino acids than in plant species-poor mixtures. Thus further research investigating diversified farming systems, including pollinator-friendly practices to reveal the attractiveness of different plant species. More diversified field margins and grasslands, for the maintenance of pollinator services for sustainable provision of crop pollination.