Ecological drivers of a vector borne pathogen: distribution and abundance of Borrelia burgdorferi sensu lato and its vector Ixodes ricinus in Scotland

Vector-borne disease emergence in recent decades has been associated with different environmental drivers including changes in habitat, hosts and climate. Lyme borreliosis is among the most important vector-borne diseases in the Northern hemisphere and is an emerging disease in Scotland. Transmitted by Ixodid tick vectors between large numbers of wild vertebrate host species, Lyme borreliosis is caused by bacteria from the Borrelia burgdorferi sensu lato species group. Ecological studies can inform how environmental factors such as host abundance and community composition, habitat and landscape heterogeneity contribute to spatial and temporal variation in risk from B. burgdorferi s.l. In this thesis a range of approaches were used to investigate the effects of vertebrate host communities and individual host species as drivers of B. burgdorferi s.l. dynamics and its tick vector Ixodes ricinus. Host species differ in reservoir competence for B. burgdorferi s.l. and as hosts for ticks. Deer are incompetent transmission hosts for B. burgdorferi s.l. but are significant hosts of all life-stages of I. ricinus. Rodents and birds are important transmission hosts of B. burgdorferi s.l. and common hosts of immature life-stages of I. ricinus. In this thesis, surveys of woodland sites revealed variable effects of deer density on B. burgdorferi prevalence, from no effect (Chapter 2) to a possible ‘dilution’ effect resulting in lower prevalence at higher deer densities (Chapter 3). An invasive species in Scotland, the grey squirrel (Sciurus carolinensis), was found to host diverse genotypes of B. burgdorferi s.l. and may act as a spill-over host for strains maintained by native host species (Chapter 4). Habitat fragmentation may alter the dynamics of B. burgdorferi s.l. via effects on the host community and host movements. In this thesis, there was lack of persistence of the rodent associated genospecies of B. burgdorferi s.l. within a naturally fragmented landscape (Chapter 3). Rodent host biology, particularly population cycles and dispersal ability are likely to affect pathogen persistence and recolonization in fragmented habitats. Heterogeneity in disease dynamics can occur spatially and temporally due to differences in the host community, habitat and climatic factors. Higher numbers of I. ricinus nymphs, and a higher probability of detecting a nymph infected with B. burgdorferi s.l., were found in areas with warmer climates estimated by growing degree days (Chapter 2). The ground vegetation type associated with the highest number of I. ricinus nymphs varied between studies in this thesis (Chapter 2 & 3) and does not appear to be a reliable predictor across large areas. B. burgdorferi s.l. prevalence and genospecies composition was highly variable for the same sites sampled in subsequent years (Chapter 2). This suggests that dynamic variables such as reservoir host densities and deer should be measured as well as more static habitat and climatic factors to understand the drivers of B. burgdorferi s.l. infection in ticks. Heterogeneity in parasite loads amongst hosts is a common finding which has implications for disease ecology and management. Using a 17-year data set for tick infestations in a wild bird community in Scotland, different effects of age and sex on tick burdens were found among four species of passerine bird (Chapter 5). There were also different rates of decline in tick burdens among bird species in response to a long term decrease in questing tick pressure over the study. Species specific patterns may be driven by differences in behaviour and immunity and highlight the importance of comparative approaches. Combining whole genome sequencing (WGS) and population genetics approaches offers a novel approach to identify ecological drivers of pathogen populations. An initial analysis of WGS from B. burgdorferi s.s. isolates sampled 16 years apart suggests that there is a signal of measurable evolution (Chapter 6). This suggests demographic analyses may be applied to understand ecological and evolutionary processes of these bacteria. This work shows how host communities, habitat and climatic factors can affect the local transmission dynamics of B. burgdorferi s.l. and the potential risk of infection to humans. Spatial and temporal heterogeneity in pathogen dynamics poses challenges for the prediction of risk. New tools such as WGS of the pathogen (Chapter 6) and blood meal analysis techniques will add power to future studies on the ecology and evolution of B. burgdorferi s.l.

Breaking old and new ground: a comparative study of coastal and inland naming in Berwickshire

This thesis investigates the place-names of four parishes in Berwickshire and compares coastal and inland naming patterns. Berwickshire is a large county that borders on northern England and historically formed part of Anglo-Saxon Northumbria. Partly due to the survival of extensive archives from the medieval priory of Coldingham, preserved in Durham Cathedral Archives, this county holds some of Scotland’s earliest recorded place- names. The parishes that form the research area are grouped together in the north-east of the county. Two of these parishes, Abbey St Bathans and Bunkle & Preston, are inland, and two, Cockburnspath and Coldingham, have extensive coastlines. The diversity of this group of parishes allows a comparative study of the place-names of coastal and inland areas to be undertaken. The topography of Berwickshire’s thirty-two parishes is very varied, and the four parishes have been chosen to reflect this range of landscapes. The place-names within the four parishes examined in this thesis derive almost exclusively from Old English, Older Scots, Modern Scots including Standard Scottish English, with a small minority derived from Old Norse, Gaelic, and Brittonic. The chronology of Old English, Older Scots, and Modern Scots is defined as given in the Concise Scots Dictionary: Old English is the period up to 1100, Older Scots is the period 1100-1700, and Modern Scots is the period 1700 onwards (CSD, 1985: xiii). Often with place-names it is not possible to give a precise dating for the coining of a toponym. For the purposes of this study, the language label given for a toponym is that of the date of the earliest record of the place-name with earlier linguistic evidence supplementing discussion. This thesis focuses on the names of topographic features, for example hills, rocks and woodland, and the role of perception in their naming. In order to compare the role of perception in inland and coastal naming, this thesis includes a diachronic study of the toponymy of the research area, along with two case studies. The first of these is a study of the toponymy of relief features, which focuses on generic elements in order to compare the perception of one type of referent in the two environments. The second is a study of the ‘colour’ category, which focuses on qualifying elements in order to compare the use of colour terms in the two environments. This thesis is the first comparative study of inland and coastal place-names, and it is one of the first to investigate new ways of using fieldwork as a central part of its methodology. In doing so it proposes innovative and nuanced ways to understand the toponymy of diverse landscapes within a community.