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    ABAtE : active bacteriophages for AFB eradication : a thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Genetics at Massey University, Auckland, New Zealand
    (Massey University, 2023) Kok, Danielle
    The European honey bee (Apis mellifera) is one of the most important livestock animals in New Zealand. Their value comes from a combination of pollination services and the production of honey for export, notably mānuka honey. American Foulbrood (AFB) is a disease of honey bee larvae and pupae and is caused by the bacterial pathogen, Paenibacillus larvae. AFB is the most serious disease that infects honey bees and is present in almost all countries where honey bees are found. AFB has been present in New Zealand since 1877 and spread to all parts of the country within 10 years. Unlike other countries, the use of antibiotics in hives infected with P. larvae is prohibited under New Zealand law and infected hives must be destroyed immediately. Bacteriophages (phages) are a well-studied alternative to antibiotics. Phages are simple viruses that kill specific bacteria and are highly abundant in the environment with an estimated 1031 globally. Phages have been shown to work effectively as a prophylactic to infection from certain diseases. With the growing antimicrobial resistance crisis, phages are becoming a well-studied and promising alternative to antibiotics. The aim of this research was to investigate the use of phages as a preventative measure against AFB. Previous work undertaken in other laboratories around the world has shown that phages can be isolated from healthy hives and nearby soil and that AFB pathogens are susceptible to destruction by these phages. In this work, we collected soil samples using citizen led science from hives throughout New Zealand. From soil samples provided we isolated 26 novel phages that are destructive to P. larvae. Selected phages were combined into a cocktail and tested against vegetative forms of P. larvae in in-lab testing. All phages were also sequenced and annotated and compared to other P. larvae phages that have been isolated around the world. This project: ABAtE (Active Bacteriophages for AFB Elimination), provides the groundwork study for an innovative approach to naturally protecting NZ beehives against AFB.
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    Microbiota in the honey bee gut and their association with bee health : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy (PhD) in Ecology at Massey University School of Agriculture and Environment, Palmerston North, New Zealand
    (Massey University, 2020) Taylor, Michelle
    European honey bees (Apis mellifera) are the most prevalent bee species globally. Honey bees play a key role in human welfare as their pollination services support both the ecological viability of wild and native plants, and the economic viability of numerous nut, fruit, and vegetable crops. A decline in unmanaged pollinators in both natural and managed ecosystems, has resulted in an increased reliance on honey bees. Despite economic globalisation and increased demand for food over the past several decades contributing to an increase in the total number of honey bee colonies worldwide, annual colony mortality is high and has been attributed to seasonal conditions, poor management practice, outbreaks of pest and disease, pesticide poisoning, and the cost of management. It has been globally hypothesised that the cause of unexplained ‘rapid’ and ‘incremental’ colony loss, may result from interactions with bee pathogens (such as Nosema spp.), environmental factors and beekeeping management. The social and foraging behaviour of bees ensures that the gut, with its bacterial residents, is the conduit for assimilating of nutrients, antibiotics and oral poison, as well as the ingress and potential reservoir for gut pathogens. Characterisation of the bacterial community within the honey bee gut may provide further insight as to how these factors may affect bee health. New Zealand honey bees have been largely bred in isolation from the rest of the world, and thus potentially developed their own gut microbiome in response to factors specific to New Zealand. These include sources of native floral nectars (e.g. mānuka and rātā), the prohibition of antibiotics for disease management, and the absence of some global honey bee gut pathogens. My research is the first to characterise the bacterial profiles and identify the relative abundance of core and less dominant bacteria in the gut of New Zealand honey bees. Diet and pathogens known to cause poor honey bee health were examined for their influence on gut bacteria. Bacterial phylotypes in the honey bee gut were identified by sequencing a fragment of the 16S rRNA gene. The problematic assignation of reliable taxonomic information for recently characterised honey bee gut bacteria was overcome by developing a customised 16S rRNA BLAST database that is compatible with QIIME2 sequencing software. This database has now been made available for other users. The five dominant core honey bee gut bacteria identified internationally were present in all apiaries/regions within New Zealand. Eight phylotypes were only identified in colonies deemed ‘sick’ by beekeepers. Three phylotypes may potentially be used as indicators of poor bee health: the family Rhizobiaceae, and the genera Serratia and Acetobacter. Although each apiary was broadly similar in bacterial composition, the environmental conditions of each apiary appeared to influence the bacterial composition, in particular the available foraging sources. In contrast to international reports, the microsporidian gut pathogen Nosema ceranae that shifted from the Asian honey bee Apis ceranae to the European honeybee in 2004 and was identified in New Zealand around 2007, does not appear to have outcompeted Nosema apis. The latter was likely brought to New Zealand with the first bees in the 19th century. In my survey N. apis was identified in all sick apiaries whereas N. ceranae was only identified in one of the sick apiaries. Comparison of the gut bacteria in New Zealand bees with those from a pilot trial conducted in Connecticut, USA demonstrates that the dominant core bacteria are internationally widespread, and suggests that they have remained stable within an isolated population for over 60 years. This highlights the importance of the symbiotic relations that these gut bacteria have with honey bees. However, nine phylotypes were present only in the New Zealand samples, suggesting that some phylotypes may have adapted to New Zealand conditions or that dysbiosis may have occurred within New Zealand or elsewhere. This is the first example in the honey bee literature of DNA being analysed using different hypervariable regions. The variation between the number of amplicon sequence variants and their relative abundances highlight the importance of comparing data extracted using similar methodologies. I observed seasonal variation in the bacterial composition by examining five hives throughout a 12-month period. Gut bacteria in summer bees were the most diverse, autumn and winter bees had lesser diversity, and spring bees had the least diversity. This suggests that the increased bee population in spring may result in a cleansing of less prevalent bacteria for the year ahead. The relative abundance of G. apicola and S. alvi did alter within individual bees throughout the year suggesting that these species may alter their abundance in response to occurrences within the gut and this may ultimately influence bee health and metabolism. The relative abundance of Rhizobiaceae peaked in winter when the bees live longer and often have elevated pathogen infections. The relative abundance of Rhizobiaceae exceeded that of all dominant core phylotypes, except Lactobacillus spp. This supports my hypothesis that Rhizobiaceae may be a useful early indicator of poor bee health. Sucrose-rich diets, often fed to bees during periods of scarce food supply, were shown to increase the relative abundances of three less dominant core bacteria; Rhizobiaceae, Acetobacteraceae, and Lactobacillus kunkeei, and decreased the relative abundance of the core species Frischella perrara. In combination, these diets significantly altered the bacterial composition. Acetogenic bacteria from the Rhizobiaceae and Acetobacteraceae families increased two- to five-fold when bees were fed sucrose, suggesting that sucrose fuels the proliferation of specific low-abundance primary sucrose-feeders. The gut pathogen N. apis did not appear to disrupt the development of Gilliamella apicola, which normally forms the outer layer of the biofilm in the luminal surface of the honey bee ileum. A gut slurry inoculation from older worker bees increased abundance of bacterial phylotypes in newly emerged workers (NEWs), thus supporting the limited literature that NEWs acquire gut bacteria from worker bees. This study also confirms that NEWs are not axenic when they emerge from their cells as their guts contain low levels of G. apicola, S. alvi, L. apis, L. mellis, Lactobacillus spp., Bifidobacterium spp., Serratia spp., Acetobacter spp., Rhizobiaceae, and Cyanobacteria. Finally, this research also identified a correlation between the lack of abundant bacteria in the honey bee gut with an increase in the opportunistic colonising bacteria Rhizobiaceae and Serratia. This is further evidence in support of my suggestion that the family Rhizobiaceae contains opportunistic bacteria and that the relative abundance of this family in honey bee guts may be a useful indicator of poor bee health. This work is thus the first study to examine gut bacteria in New Zealand honey bees and I have demonstrated that environmental factors and diet influence gut bacterial composition which may influence honey bee health and metabolism.
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    Companion biota associated with Leptospermum scoparium (mānuka; Myrtaceae) : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Ecology at Massey University, Manawatū, New Zealand
    (Massey University, 2018) Bohórquez Rodríguez de Medina, Julia
    Leptospermum scoparium (mānuka; Myrtaceae) is involved in three crucial ecological interactions that might affect nectar production, and the New Zealand honey industry. First, these plants can be affected by scale insect infestation which have the potential to affect plant health, second, they provide nectar for honey bees (mānuka honey), and third, they are hosts for, and may receive benefits from, dual mycorrhizal fungal associations (both ecto- and endo-). The understanding of these interactions is very important for the honey industry as well as for New Zealand ecosystems. However, there is limited knowledge about the influence of scale insects and mycorrhizal fungi on plant growth and nectar production, and the influence of honey bee visitation on the honey making-process. To better understand the significance of these interactions, a variety of methods, including behavioural observations, histological, molecular, and taxonomic techniques, were used in this thesis. Findings showed that the eriococcids Acanthococcus campbelli and Acanthococcus leptospermi are now the main species on L. scoparium, rather than Acanthococcus orariensis, which was the main causative agent of the mānuka blight in the 1940’s and 1960’s. Whereas the distribution of A. leptospermi was previously reported, the distribution of A. campbelli across New Zealand’s islands was illustrated for the first time in this thesis. Other scale insect species classified within the families Coelostomidiidae, Diaspididae, and Pseudococcidae were also found, but their incidence and abundance was typically lower in comparison to the family Eriococcidae. The number of eriococcids was reduced by the application of an Insect Growth Regulator (IGR) on six different cultivars in a split plot designed experiment, but cultivars differed in response to the insecticide treatment. Using the same common garden design, but just the unsprayed plants, honey bees showed a preference for the cultivar with the highest nectar sugar content and nectar DHA content. However, sugar, rather than DHA, was the best predictor of visitation pattern. The number of honey bee visits increased at midday as the day warmed up. The overall number of flowers estimated per plant was included in the model, but did not drive the visit number as, for example, it was found that the cultivar with the highest estimated number of flowers was less visited. Bioinformatics analysis revealed the association of L. scoparium with at least 25 fungal classes, including 16 ectomycorrhizal (EcM) fungal lineages and eight arbuscular mycorrhizal (AM) families. The majority of mycorrhizal fungal lineages were shared among cultivated and wild plants at the three studied sites, which suggests that cultivated plants are naturally colonised by mycorrhizal fungi. The EcM fungal lineages /cortinarius, /laccaria, /tomentella-thelephora, and the AM families Glomeraceae and Claroideoglomeraceae were the most abundant. Among the EcM fungal species, Laccaria glabripes and the endemic EcM fungal species Clavulina subrugosa, Cortinarius waiporianus and Dermocybe indotata were revealed as the most abundant. The presence of the exotic EcM fungal species Amanita muscaria was limited and mainly found in cultivated plants, that had established on a site previously with Pinus radiata. The cosmopolitan AM fungal species Rhizophagus irregularis and Claroideoglomus lamellosum were the dominant species found in both cultivated and wild plants. Among cultivated and wild plants, wild plants appeared to be colonised by a more diverse mycorrhizal fungal community. For instance, the lineage /russula-lactarius was more abundant in wild plants than in cultivated plants. The presence of /russula-lactarius and other lineages and species could be improving host performance (seed establishment, drought tolerance, pathogen resistance, and plant growth) on wild plants. However, the absence of some of the mycorrhizal fungal species from cultivated plants, which could be present on wild plants, could limit the potential yield of L. scoparium plantation. Finding suitable combinations of mycorrhizal fungal inoculum could help optimise the development of L. scoparium, nectar production, and subsequently the New Zealand mānuka honey industry.
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    Knowing the honey bee : a multispecies ethnography : a thesis presented in partial fulfilment of the requirements for the degree of Master of Arts in Social Anthropology at Massey University, Palmerston North, New Zealand
    (Massey University, 2017) Luttrell, Jordan
    Multispecies scholarship argues that the non-human has been relegated to the background of discussions about who and what inhabits and shapes the world. This thesis engages with this discussion as an experimental multispecies ethnography with honey bees in Manawatu, New Zealand. I aim to centre the honey bee in ethnography through engagement in the practice of fieldwork as well as the representation of the findings of this engagement. The honey bee is commonly known as an introduced, domesticated species, kept by humans in beehives in apiculture. This conceals the agency of the honey bee, rendering it passive, productive and compliant to the desires of humans, or in need of human intervention for survival. To view the agency of the bee I undertook embodied, performative ethnography, interviewing beekeepers and becoming one myself. My methodology, which was shaped by the bee, traced the networks that honey bees were enrolled in. Encounters were awkward, one-sided, and sometimes dangerous. The representation of honey bees demands an approach which attends to multiple, distinct accounts of honey bee worlds, because the bee is a lively agent, contributing to, experiencing, and communicating about the multiple networks in which it is engaged. As such, the findings of this thesis are presented in three accounts of encounters with honey bees. These accounts are distinct, capturing the honey bee in different networks, but are also distinct in their narrative styles, progressing from a description of honey networks in the spirit of Actor-Networks, to writing with honey bee narrator in poetry. Ethnographic representation is inevitably partial and an act of imagination. However, becoming sensitive to the ‘bee-ness’ of the bee; the waggle, hum and sting, and employing narrative inspired by the multisensory apiary, in other words, shaping representation with honey bees in mind, is an act of privileging honey bees in writing, and exploring what more can be said of, and with, the bee.
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    The impact of honey bees on montane ecosystems within Tongariro National Park : a thesis presented in partial fulfilment of the requirements for the degree of Master of Science in Ecology at Massey University
    (Massey University, 1996) Murphy, Claire
    A study of the effects of honey bees on montane ecosystems was conducted during the summers of 1993/1994 and 1994/1995 at Tongariro National Park. Three possible effects of the introduced honey bee were examined. The primary aims of the study were to identity areas with and without honey bees and to identify differences in the pollination success of a weed species, heather (Calluna vugaris) and a native species, the New Zealand flax (Phormium tenax) under different pollinator regimes, and to examine differences in the composition of native pollinator communities in these different bee areas. The impact of honey bees on the reproductive success of heather, an important weed species in Tongariro National Park was examined over two flowering seasons. Insect visitation rates on heather flowers were low at each of the four study sites. Bagging plants to exclude insect flower visitors had little effect on female fitness. The potential of other pollen vectors, wind and thrips, as pollinators of heather was also examined. Both were determined to have a negative effect on several measures of female success, including pollen deposition, pollen tube formation, and pollination levels. However it appears that none of the pollen vectors (honey bees, wind or thrips) significantly effect the overall fitness of heather in terms of the viable seed produced. The second part of the study examined the impact of honey bees on the pollination systems of a native plant species. Flax is thought to be predominantly bird pollinated, however, the floral resources are also utilised by a variety of native and introduced insect species. At some sites birds were either not present or rarely used the flowers. Seed set in flax was highest in heavily bird pollinated sites. The results also suggest, however, that flax has a flexible pollinating system that enables it to maintain a range of fruit and seed set levels under the different pollintor regimes. The abundance and diversity of insect flower visitors on manuka and Hebe stricta, two common subalpine shrubs, was highly variable between sites, and between observation periods. Some of this variation may be ascribed to differences in the weather or to altitude. However, I have shown that the abundance and diversity of diptera appears to be strongly influenced by levels of honey bee activity. This indicates that honey bees do play a role in determining the structure of pollinator communities and may be displacing a significant component of the native pollinating fauna.