Gavin J. Wilson

A review of methods to estimate the abundance of terrestrial carnivores using field signs and observation

This paper reviews field methods for estimating and monitoring the abundance of terrestrial carnivores that do not involve capture. Effective methods of monitoring abundance are important tools for the management and conservation of many species. The development of methods for carnivores presents particular challenges, as they are often secretive and widely dispersed. Nevertheless, a variety of approaches based on direct observations and quantification of field signs have been employed. These techniques are described in relation to carnivore ecology and resource implications, and the advantages and deficiencies of each are discussed with reference to case studies. Estindasvore G.ls J W033 Ga J. Rlah

Reliable microsatellite genotyping of the Eurasian badger ( Meles meles ) using faecal DNA

The potential link between badgers and bovine tuberculosis has made it vital to develop accurate techniques to census badgers. Here we investigate the potential of using genetic profiles obtained from faecal DNA as a basis for population size estimation. After trialling several methods we obtained a high amplification success rate (89%) by storing faeces in 70% ethanol and using the guanidine thiocyanate/silica method for extraction. Using 70% ethanol as a storage agent had the advantage of it being an antiseptic. In order to obtain reliable genotypes with fewer amplification reactions than the standard multiple‐tubes approach, we devised a comparative approach in which genetic profiles were compared and replication directed at similar, but not identical, genotypes. This modified method achieved a reduction in polymerase chain reactions comparable with the maximum‐likelihood model when just using reliability criteria, and was slightly better when using reliability criteria with the additional proviso that alleles must be observed twice to be considered reliable. Our comparative approach would be best suited for studies that include multiple faeces from each individual. We utilized our approach in a well‐studied population of badgers from which individuals had been sampled and reliable genotypes obtained. In a study of 53 faeces sampled from three social groups over 10 days, we found that direct enumeration could not be used to estimate population size, but that the application of mark–recapture models has the potential to provide more accurate results.

Bacillus Calmette-Guérin vaccination reduces the severity and progression of tuberculosis in badgers

Control of bovine tuberculosis (TB) in cattle has proven particularly challenging where reservoirs of infection exist in wildlife populations. In Britain and Ireland, control is hampered by a reservoir of infection in Eurasian badgers (Meles meles). Badger culling has positive and negative effects on bovine TB in cattle and is difficult, costly and controversial. Here we show that Bacillus Calmette-Guérin (BCG) vaccination of captive badgers reduced the progression, severity and excretion of Mycobacterium bovis infection after experimental challenge. In a clinical field study, BCG vaccination of free-living badgers reduced the incidence of positive serological test results by 73.8 per cent. In common with other species, BCG did not appear to prevent infection of badgers subjected to experimental challenge, but did significantly reduce the overall disease burden. BCG vaccination of badgers could comprise an important component of a comprehensive programme of measures to control bovine TB in cattle.

Mating system of the Eurasian badger, Meles meles, in a high density population

Badgers are facultatively social, forming large groups at high density. Group‐living appears to have high reproductive costs for females, and may lead to increased levels of inbreeding. The extent of female competition for reproduction has been estimated from field data, but knowledge of male reproductive success and the extent of extra‐group paternity remains limited. Combining field data with genetic data (16 microsatellite loci), we studied the mating system of 10 badger social groups across 14 years in a high‐density population. From 923 badgers, including 425 cubs, we were able to assign maternity to 307 cubs, with both parents assigned to 199 cubs (47%) with 80% confidence, and 14% with 95% confidence. Age had a significant effect on the probability of reproduction, seemingly as a result of a deficit of individuals aged two years and greater than eight years attaining parentage. We estimate that approximately 30% of the female population successfully reproduced in any given year, with a similar proportion of the male population gaining paternity across the same area. While it was known there was a cost to female reproduction in high density populations, it appears that males suffer similar, but not greater, costs. Roughly half of assigned paternity was attributed to extra‐group males, the majority of which were from neighbouring social groups. Few successful matings occurred between individuals born in the same social group (22%). The high rate of extra‐group mating, previously unquantified, may help reduce inbreeding, potentially making philopatry a less costly strategy.

Is Mycobacterium bovis in the environment important for the persistence of bovine tuberculosis

Mycobacterium bovis is the causative agent of bovine tuberculosis (bTB) in cattle and wildlife. Direct aerosol contact is thought to be the primary route of infection between conspecifics, whereas indirect transmission via an environmental reservoir of M. bovis is generally perceived not to be a significant source for infection. Here, we report on the application of molecular technology (PCR) to quantify the prevalence of M. bovis in the environment and to explore its epidemiological significance. We show that the detectability of viable M. bovis at badger setts and latrines is strongly linked to the frequency of M. bovis excretion by infected badgers, and that putative M. bovis in the environment is prevalent on a large proportion of endemic cattle farms in Britain. These results raise important questions about the role of an environmental reservoir in bTB persistence.

Using isolation-by-distance-based approaches to assess the barrier effect of linear landscape elements on badger (Meles meles) dispersal

As the European badger (Meles meles) can be of conservation or management concern, it is important to have a good understanding of the species’ dispersal ability. In particular, knowledge of landscape elements that affect dispersal can contribute to devising effective management strategies. However, the standard approach of using Bayesian clustering methods to correlate genetic discontinuities with landscape elements cannot easily be applied to this problem, as badger populations are often characterized by a strong confounding isolation‐by‐distance (IBD) pattern. We therefore developed a two‐step method that compares the location of pairs of related badgers relative to a putative barrier and utilizes the expected spatial genetic structure characterized by IBD as a null model to test for the presence of a barrier. If a linear feature disrupts dispersal, the IBD pattern characterising pairs of individuals located on different sides of a putative barrier should differ significantly from the pattern obtained with pairs of individuals located on the same side. We used our new approach to assess the impact of rivers and roads of different sizes on badger dispersal in western England. We show that a large, wide river represented a barrier to badger dispersal and found evidence that a motorway may also restrict badger movement. Conversely, we did not find any evidence for small rivers and roads interfering with badger movement. One of the advantages of our approach is that potentially it can detect features that disrupt gene flow locally, without necessarily creating distinct identifiable genetic units.

Genetic evidence that culling increases badger movement: implications for the spread of bovine tuberculosis

The Eurasian badger (Meles meles) has been implicated in the transmission of bovine tuberculosis (TB, caused by Mycobacterium bovis) to cattle. However, evidence suggests that attempts to reduce the spread of TB among cattle in Britain by culling badgers have mixed effects. A large‐scale field experiment (the randomized badger culling trial, RBCT) showed that widespread proactive badger culling reduced the incidence of TB in cattle within culled areas but that TB incidence increased in adjoining areas. Additionally, localized reactive badger culling increased the incidence of TB in cattle. It has been suggested that culling‐induced perturbation of badger social structure may increase individual movements and elevate the risk of disease transmission between badgers and cattle. Field studies support this hypothesis, by demonstrating increases in badger group ranges and the prevalence of TB infection in badgers following culling. However, more evidence on the effect of culling on badger movements is needed in order to predict the epidemiological consequences of this control strategy. Here, analysis of the genetic signatures of badger populations in the RBCT revealed increased dispersal following culling. While standard tests provided evidence for greater dispersal after culling, a novel method indicated that this was due to medium‐ and long‐distance dispersal, in addition to previously reported increases in home‐range size. Our results also indicated that, on average, badgers infected with M. bovis moved significantly farther than did uninfected badgers. A disease control strategy that included culling would need to take account of the potentially negative epidemiological consequences of increased badger dispersal.

Experimental evidence of competitive release in sympatric carnivores

Changes in the relative abundance of sympatric carnivores can have far-reaching ecological consequences, including the precipitation of trophic cascades and species declines. While such observations are compelling, experimental evaluations of interactions among carnivores remain scarce and are both logistically and ethically challenging. Carnivores are nonetheless a particular focus of management practices owing to their roles as predators of livestock and as vectors and reservoirs of zoonotic diseases. Here, we provide evidence from a replicated and controlled experiment that culling Eurasian badgers Meles meles for disease control was associated with increases in red fox Vulpes vulpes densities of 1.6–2.3 foxes km−2. This unique experiment demonstrates the importance of intraguild relations in determining species abundance and of assessing the wider consequences of intervention in predator populations.

Vaccinating badgers (Meles meles) against Mycobacterium bovis: the ecological considerations.

Bovine tuberculosis (TB) is a serious zoonotic disease, which despite a largely successful test and slaughter programme has persisted in cattle herds in parts of the UK. The badger (Meles meles) is widely considered to represent a significant wildlife reservoir for the transmission of Mycobacterium bovis to cattle, and has been the subject of a variety of culling strategies since the mid 1970s. Nevertheless, the incidence of herd breakdowns has continued to rise, and the efficacy of culling is currently the subject of a large-scale field trial. One potential alternative tool for the management of disease in wildlife populations is vaccination. However, the successful development of an effective vaccine and a strategy for its delivery will require careful consideration of the practical constraints imposed by ecological factors. In the current paper, we discuss relevant ecological and epidemiological characteristics of badger populations and practical aspects of vaccine delivery in the field.

An evaluation of distance sampling to estimate badger (Meles meles) abundance

Distance sampling is a common and increasingly used method of estimating animal abundance in conservation and management programmes. The precision of a distance sampling survey can be estimated from the field data, but accuracy can only be evaluated by comparison with the true population size. For wild mammal populations, such opportunities are rare. The high-density badger Meles meles population at Woodchester Park in Gloucestershire, U.K. has been routinely and intensively live-trapped for the past 30 years. The estimates of abundance based on mark–recapture analyses of the live-trapping data provided a reliable baseline against which to evaluate distance sampling for a badger population. A distance sampling survey was carried out in autumn 2002 by using spotlighting along line transects. Owing to the impractical nature of spotlighting in closed habitats, such as woodland, surveys were conducted in open habitat only. A radio-tracking study provided behavioural data that were used to estimate the proportion of the population available for sampling in open habitats. This was incorporated as a multiplier into the DISTANCE analysis. The estimate of abundance from distance sampling was 76 badgers (% CV=42.2), using a multiplier based on radio-tracking data from autumn only. When the multiplier was derived from radio-tracking data from across the whole year, based on a larger sample of tracked badgers, the distance sampling abundance estimate was 77 badgers (% CV=25.9). These compared favourably to the mark–recapture estimate of 68 badgers (% C.V=7.4). The lack of precision in the distance sampling density estimate was largely owing to the inclusion of the multiplier. Distance sampling combined with data on habitat use can produce reliable estimates of badger abundance in areas of medium to high density, particularly in landscapes of primarily open habitat.