Density and abundance of badger social groups in England and Wales in 2011-2013

This is the final version of the article. Available from Nature Publishing Group via the DOI in this record.In the United Kingdom, European badgers Meles meles are a protected species and an important wildlife reservoir of bovine tuberculosis. We conducted a survey of badger dens (main setts) in 1614 1 km squares across England and Wales, between November 2011 and March 2013. Using main setts as a proxy for badger social groups, the estimated mean density of badger social groups in England and Wales was 0.485 km(-2) (95% confidence interval 0.449-0.521) and the estimated abundance of social groups was 71,600 (66,400-76,900). In the 25 years since the first survey in 1985-88, the annual rate of increase in the estimated number of badger social groups was 2.6% (2.2-2.9%), equating to an 88% (70-105%) increase across England and Wales. In England, we estimate there has been an increase of 103% (83-123%) in badger social groups, while in Wales there has been little change (-25 to +49%).We are grateful to the thousands of landowners for their kind co-operation in granting access to their land. This study was funded by the Department for Environment, Food and Rural Affairs, as part of England and Wales national research activities. Fieldwork was conducted by staff of the National Wildlife Management Centre. Access to data from the 1985–88 survey was licensed by the Joint Nature Conservation Committee, to whom the rights of the Nature Conservancy Council had passed

Contact chains of cattle farms in Great Britain

This is the final version. Available from the publisher via the DOI in this record.Further information and figures supporting this article have been uploaded as part of the electronic supplementary material. Underlying data consist of every movement of cattle between all farms in Great Britain. Aside from the size of the dataset, there are substantial issues of confidentiality (locations, trading practices) and commercial sensitivity in these data. They are collated and managed by Defra, via the Animal and Plant Health Agency, who grant access to the data with specific permissions for specific studies. In practice, this means that the data can be used for the stated purpose only, and making the data publicly accessible would not conform to the licence the authors have been granted to use these data. With the agreement of the journal’s Editorial Office, the authors will not be able to make the dataset available on this occasion, but encourage readers, referees and editors to contact the Animal and Plant Health Agency data manager for data access requests. At the time of submission, the data manager is Andy Mitchell ([email protected])Network analyses can assist in predicting the course of epidemics. Time-directed paths or ‘contact chains’ provide a measure of hostconnectedness across specified timeframes, and so represent potential pathways for spread of infections with different epidemiological characteristics. We analysed networks and contact chains of cattle farms in Great Britain using Cattle Tracing System data from 2001 to 2015. We focused on the potential for between-farm transmission of bovine tuberculosis, a chronic infection with potential for hidden spread through the network. Networks were characterized by scale-free type properties, where individual farms were found to be influential ‘hubs’ in the network. We found a markedly bimodal distribution of farms with either small or very large ingoing and outgoing contact chains (ICCs and OCCs). As a result of their cattle purchases within 12-month periods, 47% of British farms were connected by ICCs to more than 1000 other farms and 16% were connected to more than 10 000 other farms. As a result of their cattle sales within 12-month periods, 66% of farms had OCCs that reached more than 1000 other farms and 15% reached more than 10 000 other farms. Over 19 000 farms had both ICCs and OCCs reaching more than 10 000 farms for two or more years. While farms with more contacts in their ICCs or OCCs might play an important role in disease spread, farms with extensive ICCs and OCCs might be particularly important by being at higher risk of both acquiring and disseminatinginfections.Biotechnology and Biological Science Research Council (BBSRC

Abundance of badgers (Meles meles) in England and Wales

This is the author accepted manuscript. The final version is available on open accessfrom the publisher via the DOI in this recordThe European badger (Meles meles) is of considerable interest in the UK as it is both a protected species and the main wildlife reservoir for bovine tuberculosis infection in cattle. While there have been three national badger surveys in the 1980s, 1990s and 2011-13, using the number of badger main setts as a proxy for the abundance of badger social groups, none has combined contemporary data on social group size at landscape and national scales. We estimated social group size by genotyping hair samples collected at 120 main setts across England and Wales and employing a capture-mark-recapture method based on genotypes. The estimated mean social group size in England and Wales was 6.74 (±0.63) badgers. There was considerable variation in badger social group size among Land Class Groups (LCGs), with a low of 2.67 in LCG3 and a high of 7.92 in LCG4. Combining these results with the recent Badger Sett Survey of England and Wales, we estimate there are approximately 485,000 badgers (95% confidence intervals 391,000-581,000) in England and Wales. Although direct comparison with previous estimates is not ideal owing to methodological differences, our results are consistent with a marked increase in the badger population of England and Wales since the 1980s.This study was funded by the Department for the Environment, Food and Rural Affairs, as part of their England and Wales national research activities

Characterization of potential superspreader farms for bovine tuberculosis: A review

This is the final version. Available on open access from Wiley via the DOI in this recordBACKGROUND: Variation in host attributes that influence their contact rates and infectiousness can lead some individuals to make disproportionate contributions to the spread of infections. Understanding the roles of such 'superspreaders' can be crucial in deciding where to direct disease surveillance and controls to greatest effect. In the epidemiology of bovine tuberculosis (bTB) in Great Britain, it has been suggested that a minority of cattle farms or herds might make disproportionate contributions to the spread of Mycobacterium bovis, and hence might be considered 'superspreader farms'. OBJECTIVES AND METHODS: We review the literature to identify the characteristics of farms that have the potential to contribute to exceptional values in the three main components of the farm reproductive number - Rf : contact rate, infectiousness and duration of infectiousness, and therefore might characterize potential superspreader farms for bovine tuberculosis in Great Britain. RESULTS: Farms exhibit marked heterogeneity in contact rates arising from between-farm trading of cattle. A minority of farms act as trading hubs that greatly augment connections within cattle trading networks. Herd infectiousness might be increased by high within-herd transmission or the presence of supershedding individuals, or infectiousness might be prolonged due to undetected infections or by repeated local transmission, via wildlife or fomites. CONCLUSIONS: Targeting control methods on putative superspreader farms might yield disproportionate benefits in controlling endemic bovine tuberculosis in Great Britain. However, real-time identification of any such farms, and integration of controls with industry practices, present analytical, operational and policy challenges.Biotechnology and Biological Sciences Research Council (BBSRC)Animal and Plant Health Agenc

Bait uptake by wild badgers and its implications for oral vaccination against tuberculosis

This is the final version. Available from Public Library of Science via the DOI in this record.Data Availability: All relevant data are within the paper and its Supporting Information files.The deployment of baits containing vaccines or toxins has been used successfully in the management of wildlife populations, including for disease control. Optimisation of deployment strategies seeks to maximise uptake by the targeted population whilst ensuring cost-effectiveness. Tuberculosis (TB) caused by infection with Mycobacterium bovis affects a broad range of mammalian hosts across the globe, including cattle, wildlife and humans. The control of TB in cattle in the UK and Republic of Ireland is hampered by persistent infection in European badgers (Meles meles). The present study aimed to determine the best strategy for maximising uptake of an oral vaccine by wild badgers, using a surrogate novel bait deployed at 40 badger social groups. Baits contained a blood-borne biomarker (Iophenoxic Acid, IPA) in order to measure consumption in badgers subsequently cage trapped at targeted setts. Evidence for the consumption of bait was found in 83% (199/240) of captured badgers. The probability that badgers had consumed at least one bait (IPA >10 μg ml-1) was significantly higher following deployment in spring than in summer. Lower uptake amongst social groups where more badgers were captured, suggested competition for baits. The probability of bait consumption was significantly higher at groups where main and outlier setts were provided with baits than at those where outliers were present but not baited. Badgers captured 10–14 days post bait feeding had significantly higher levels of bait uptake compared to those caught 24–28 days later. Uptake rates did not vary significantly in relation to badger age and whether bait was placed above ground or down setts. This study suggests that high levels of bait uptake can be achieved in wild badger populations and identifies factors influencing the potential success of different deployment strategies. The implications for the development of an oral badger vaccine are discussed.Natural Environment Research Council (NERC)Animal and Plant Health Agency (APHA

The application of statistical network models in disease research

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.Host social structure is fundamental to how infections spread and persist, and so the statistical modelling of static and dynamic social networks provides an invaluable tool to parameterise realistic epidemiological models. We present a practical guide to the application of network modelling frameworks for hypothesis testing related to social interactions and epidemiology, illustrating some approaches with worked examples using data from a population of wild European badgers Meles meles naturally infected with bovine tuberculosis. Different empirical network datasets generate particular statistical issues related to non-independence and sampling constraints. We therefore discuss the strengths and weaknesses of modelling approaches for different types of network data and for answering different questions relating to disease transmission. We argue that statistical modelling frameworks designed specifically for network analysis offer great potential in directly relating network structure to infection. They have the potential to be powerful tools in analysing empirical contact data used in epidemiological studies, but remain untested for use in networks of spatio-temporal associations. As a result, we argue that developments in the statistical analysis of empirical contact data are critical given the ready availability of dynamic network data from bio-logging studies. Furthermore, we encourage improved integration of statistical network approaches into epidemiological research to facilitate the generation of novel modelling frameworks and help extend our understanding of disease transmission in natural populations.M.J.S. is funded by a NERC standard grant (NE/M004546/1) awarded to R.A.M., D.P.C., D.J.H. and M.B., with the APHA team at Woodchester Park, UK (lead scientist is R.J.D.) as project partners

Demographic buffering and compensatory recruitment promotes the persistence of disease in a wildlife population.

Published onlineLETTERDemographic buffering allows populations to persist by compensating for fluctuations in vital rates, including disease-induced mortality. Using long-term data on a badger (Meles meles Linnaeus, 1758) population naturally infected with Mycobacterium bovis, we built an integrated population model to quantify impacts of disease, density and environmental drivers on survival and recruitment. Badgers exhibit a slow life-history strategy, having high rates of adult survival with low variance, and low but variable rates of recruitment. Recruitment exhibited strong negative density-dependence, but was not influenced by disease, while adult survival was density independent but declined with increasing prevalence of diseased individuals. Given that reproductive success is not depressed by disease prevalence, density-dependent recruitment of cubs is likely to compensate for disease-induced mortality. This combination of slow life history and compensatory recruitment promotes the persistence of a naturally infected badger population and helps to explain the badger's role as a persistent reservoir of M. bovis.NERCUK Department of Environment, Food and Rural Affair

Model of Selective and Non-Selective Management of Badgers (Meles meles) to Control Bovine Tuberculosis in Badgers and Cattle.

This is the author accepted manuscript. The final version is available from Public Library of Science via the DOI in this record.Bovine tuberculosis (bTB) causes substantial economic losses to cattle farmers and taxpayers in the British Isles. Disease management in cattle is complicated by the role of the European badger (Meles meles) as a host of the infection. Proactive, non-selective culling of badgers can reduce the incidence of disease in cattle but may also have negative effects in the area surrounding culls that have been associated with social perturbation of badger populations. The selective removal of infected badgers would, in principle, reduce the number culled, but the effects of selective culling on social perturbation and disease outcomes are unclear. We used an established model to simulate non-selective badger culling, non-selective badger vaccination and a selective trap and vaccinate or remove (TVR) approach to badger management in two distinct areas: South West England and Northern Ireland. TVR was simulated with and without social perturbation in effect. The lower badger density in Northern Ireland caused no qualitative change in the effect of management strategies on badgers, although the absolute number of infected badgers was lower in all cases. However, probably due to differing herd density in Northern Ireland, the simulated badger management strategies caused greater variation in subsequent cattle bTB incidence. Selective culling in the model reduced the number of badgers killed by about 83% but this only led to an overall benefit for cattle TB incidence if there was no social perturbation of badgers. We conclude that the likely benefit of selective culling will be dependent on the social responses of badgers to intervention but that other population factors including badger and cattle density had little effect on the relative benefits of selective culling compared to other methods, and that this may also be the case for disease management in other wild host populations.Funding organisations were: 1. Department for Food, Environment and Rural Affairs, and 2. Department of Agriculture and Rural Development Northern Ireland. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript

Seasonal variation in daily patterns of social contacts in the European badger Meles meles

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.Social interactions among hosts influence the persistence and spread of infectious pathogens. Daily 20 and seasonal variation in the frequency and type of social interactions will play an important role in 21 disease epidemiology, and alongside other factors may have an influence on wider disease dynamics 22 by causing seasonal forcing of infection, especially if the seasonal variation experienced by a 23 population is considerable. We explored temporal variation in within-group contacts in a high-24 density population of European badgers Meles meles naturally-infected with bovine tuberculosis. 25 Summer contacts were more likely and of longer duration during the daytime, while the frequency 26 and duration of winter contacts did not differ between day and night. In spring and autumn within-27 group contacts peaked at dawn and dusk, corresponding with when they were of shortest duration 28 with reduced potential for aerosol transmission of pathogens. Summer and winter could be critical 29 for bovine tuberculosis transmission in badgers, due to the high frequency and duration of contacts 30 during resting periods, and we discuss the links between this result and empirical data. This study 31 reveals clear seasonality in daily patterns of contact frequency and duration in species living in stable 32 social groups, suggesting that changes in social contacts could drive seasonal forcing of infection in 33 wildlife populations even when the number of individuals interacting remains similar.MJS is funded by NERC grant NE/M004546/1 awarded to RAM, DPC, DJH and MB, with RJD and the 386 APHA team at Woodchester Park, UK as project partners. Data were collected for NW’s PhD, funded 387 by Defra. We thank Jared Wilson-Aggarwal and two anonymous reviewers for useful comments and 388 Keith Silk for providing the photograph for Figure 1

Investigation into the genetic diversity in toll-like receptors 2 and 4 in the European badger Meles meles

The Toll-like receptor (TLR) genes are a conserved family of genes central to the innate immune response to pathogen infection. They encode receptor proteins, recognise pathogen associated molecular patterns (PAMPs) and trigger initial immune responses. In some host-pathogen systems, it is reported that genetic differences, such as single nucleotide polymorphisms (SNPs), associate with disease resistance or susceptibility. Little is known about TLR gene diversity in the European badger (Meles meles). We collected DNA from UK badgers, carried out PCR amplification of the badger TLR2 gene and exon 3 of TLR4 and determined DNA sequences for individual badgers for TLR2 (n=61) and TLR4 exon 3 (n=59). No polymorphism was observed in TLR4. Three TLR2 amino acid haplotype variants were found. Ninety five percent of badgers were homozygous for one common haplotype (H1), the remaining three badgers had genotypes H1/H3, H1/H2 and H2/H2. By broad comparison with other species, diversity in TLR genes in badgers seems low. This could be due to a relatively localised sampling or inherent low genetic diversity. Further studies are required to assess the generality of the low observed diversity and the relevance to the immunological status of badgers