Seasonality of reproduction in the European badger Meles meles in south-west England

The reproductive cycle of male and female badgers in the south-west of England was studied by post-mortem examination of 1875 badgers collected in 1973–80. The sample was obtained by several methods and showed that animals obtained as road casualties were not representative of the total population samples. Active spermatogenesis was present throughout the year, and in some males aged 8–9 months. A few females first ovulated as yearlings but most primiparous females were adults. Births were in general confined to the early part of the year, but ovulations and matings also occurred at other times. Almost all (97%) adult females possessed corpora lutea. Unimplanted blastocysts were found in females for 11 months of the year (over 80% of adult females in July-November). Implanted embryos were found in only 32% of adults in January and February: the number of blastocysts per female and the number of implanted embryos per female were not significantly different, indicating that sows implanted all blastocysts or none. Foetal mortality was 36% and post-natal losses were estimated at 42% from the proportion of lactating females.     

The role of setts in badger (Meles meles) group size, breeding success and status of TB (Mycobacterium bovis)

This paper examines the relationship between the number of occupied setts in a badger social group territory and badger group size, breeding success, and status of infection with Mycobacterium bovis (TB). The data used were from a long-term epidemiological and ecological study of a high-density population of badgers Meles meles in south-west England. The number of occupied setts in a social group was significantly and positively related to the number of badgers caught in the social group, so that as a social group increases in size, badgers occupy more of the available setts. This relationship remained significant when numbers of adults, adult males and adult females were examined. The number of breeding females, number of cubs and sex ratio was not related to the number of occupied setts in a social group. It is possible that the advantages to breeding females of a larger number of setts available to breed in might be outweighed by the increased aggression found in larger groups. The TB score for prevalence and for incidence of social groups was significantly and positively related to the number of occupied setts in a social group, such that the more occupied setts there were in a territory, the higher the TB index of the group. Possibly the setts themselves contribute to the persistence of TB within social groups, or badgers infected with TB might show a difference in behaviour from uninfected badgers resulting in their increased use of outlying setts.     

Evaluating seasonal bait delivery to badgers using rhodamine B

In the UK and Ireland, research on the control of bovine tuberculosis in badgers includes the development of a palatable bait for oral delivery of a vaccine and a means of its deployment in the field. In the present study, we carried out field deployment of bait according to the established method of bait marking in early spring and early summer to compare the effects of seasonality on bait uptake rates. All baits contained rhodamine B (RhB) which was subsequently detected in the hair and whiskers of captured badgers. During the 8 days of bait feeding at 14 badger setts, 99% of baits deployed in spring, and 100% of those deployed in summer were removed. The presence of RhB in captured badgers indicated high rates of uptake amongst adult badgers in spring (93%) and summer (98%). Only cubs captured in summer showed evidence of having taken bait (91%). Between 67% and 100% of each social group was estimated to have taken bait. The detection of RhB in 96% of badgers captured at outlier setts, where bait was not fed, suggested that deployment at main setts alone may be sufficient to target a relatively high proportion of the badger population. The number of baits deployed per marked badger suggested that a similar level of uptake might be achievable using fewer baits. The results clearly demonstrate the potential value of the bait-marking methodology for delivering vaccine baits to badgers during spring and summer, but that deployment in early summer is necessary to target cubs.     

Winter hibernation and body temperature fluctuation in the Japanese badger, Meles meles anakuma

This study examined seasonal changes in body weight, hibernation period, and body temperature of the Japanese badger (Meles meles anakuma) from 1997 to 2001. Adult badgers showed seasonal changes in body weight. Between mid-December and February, badger activity almost ceased, as the animals remained in their setts most of the time. Adult male badgers were solitary hibernators; adult females hibernated either alone or with their cubs and/or yearlings. The total hibernation period of Japanese badgers ranged from 42 to 80 days, with a mean length of 60.1 days. Japanese badgers did not always spend the winters in the same sett, although they seldom changed setts during hibernation. I equipped a male cub with an intraperitoneally implanted data logger to record its body temperature between November and April, while the cub hibernated with its mother. Over the winter, the body weight of the cub decreased from 5.3 kg to 3.6 kg, a weight loss of 32.1%, and its body temperature ranged from 32.0 to 39.8 degrees C. The mean monthly body temperature was 35.1 degrees C in December, 34.8 degrees C in January, 35.9 degrees C in February, 37.1 degrees C in March, and 37.4 degrees C in April, so the monthly decrease in body temperature of this cub was not great. The results indicate that during hibernation, when body temperature is low, there is likely considerable economy of energy and a reduced demand for adipose reserves.     

Distribution and population density of badgers Meles meles in Luxembourg

1. The distribution and density of Eurasian badgers Meles meles in Luxembourg was estimated by gathering information about the location of badger setts with a questionnaire survey, by visiting 708 setts in order to classify them as ‘main setts’ or ‘outliers’, and by estimating social group size by directly counting emerging badgers. 2. Badgers were found to be widely distributed in Luxembourg, with a minimum main sett density of 0.17 setts/km². Setts were sited preferentially in forest habitat. The mean minimum group size was 4.6 badgers. 3. The Luxembourg badger population was conservatively estimated to contain at least 2010 adult and young badgers (95% CI 1674–2347) in spring 2002, equivalent to a density of 0.78 adult and young badgers/km² (95% CI 0.65–0.91). This is moderate compared to most of continental Europe.     

Use of human buildings by Eurasian badgers in the Moravskoslezské Beskydy Mountains,Czech Republic

An inspection of human buildings used by Eurasian badgersMeles meles (Linnaeus, 1758) in 28 sites in the Moravskoslezské Beskydy Mountains, Czech Republic, was carried out in 2001. The buildings inhabited or visited by badgers were as follows: wooden barns (18 cases), masonry buildings used for residential purposes (4), abandoned buildings (1), wooden sheds (2), wooden beehouses (2) and a non-residential part of a house (1). In three sites, female badgers with their cubs inhabited buildings. Badgers use the buildings more frequently in winter than in summer. Use of human buildings and the occurrence of badgers in setts in the wild in these mountains was observed in detail on a study area of 950 ha around the village of Lubno. In total, 12 setts were discovered. Eight of them were in the wild: two setts were located closer than 50 m, five between 100 m and 300 m, and one 700 m from human buildings. In four sites badgers inhabited human buildings.     

Seasonal and spatial pattern of shelter use by badgers<Emphasis Type="Italic">Meles meles</Emphasis> in Białowieża Primeval Forest (Poland)

In 1997–2001, we investigated the use of day-time shelters by radio-collared badgersMeles meles (Linnaeus, 1758) in the Białowieża Primeval Forest, eastern Poland. Each social group of badgers utilised, on average, 9 different shelters per territory (range: 4–20). The main setts, occupied for breeding and winter sleep, were also most frequently used for day-time rest throughout the year (73% of days). Badgers living in the pristine oldgrowth stands utilised larger number of shelters and spent more days in hollow trees (mainly limeTilia cordata), compared to badgers inhabiting younger secondary tree stands. Number of shelters used by individuals varied between seasons and depended on sex and age of animals. In summer, badgers used more shelters than in spring and autumn. In winter, they stayed in their main setts only. Adult males occupied more shelters and spent fewer days in the main sett than other badgers. In spring, females rearing young used only the main setts. The average underground space used by badgers within the main sett was 128 m². It was largest in summer and smallest in winter, and also varied between males and females. We proposed that, in a low-density population, badgers used several setts and other daily shelters to reduce energy expenditure when exploring their large territories and foraging. Furthermore, setts may play a role of marking sites. Analysis of the biogeographical pattern of sett use by European badgers showed that the number of setts used by social groups increased with increasing territory size, whereas the density of setts (n setts/km²) was negatively correlated with territory size. We proposed that different factors could shape the utilisation of setts by badgers in low- and high-density populations.     

Reproductive Biology Including Evidence for Superfetation in the European Badger Meles meles (Carnivora: Mustelidae)

The reproductive biology of the European badger (Meles meles) is of wide interest because it is one of the few mammal species that show delayed implantation and one of only five which are suggested to show superfetation as a reproductive strategy. This study aimed to describe the reproductive biology of female Irish badgers with a view to increasing our understanding of the process of delayed implantation and superfetation. We carried out a detailed histological examination of the reproductive tract of 264 female badgers taken from sites across 20 of the 26 counties in the Republic of Ireland. The key results show evidence of multiple blastocysts at different stages of development present simultaneously in the same female, supporting the view that superfetation is relatively common in this population of badgers. In addition we present strong evidence that the breeding rate in Irish badgers is limited by failure to conceive, rather than failure at any other stages of the breeding cycle. We show few effects of age on breeding success, suggesting no breeding suppression by adult females in this population. The study sheds new light on this unusual breeding strategy of delayed implantation and superfetation, and highlights a number of significant differences between the reproductive biology of female Irish badgers and those of Great Britain and Swedish populations.     

Seasonal and inter-individual variation in testosterone levels in badgers <Emphasis Type="Italic">Meles meles</Emphasis>: evidence for the existence of two endocrinological phenotypes

Elevated testosterone levels can lower condition and increase parasites. We analysed testosterone in 84 blood samples of wild European badgers Meles meles collected at regular intervals (winter = mating season; spring = end of mating season; summer = minor mating peak; autumn = reproductive quiescence), and related variation to body condition, subcaudal gland secretion, parasite burden, and bite wounding. All males showed elevated levels in winter and low levels in autumn. In neither season did testosterone correlate with fitness-related parameters. However, two different endocrinological phenotypes existed in spring and summer. Whilst some males lowered their testosterone to levels comparable to autumnal quiescence (Type 1), others maintained elevated levels comparable to those during winter (Type 2). In spring and summer high levels were correlated with lower body condition and increased parasite burden, and Type 2 males tended to suffer higher mortality rates than Type 1. No animals older than 6 years adopted phenotype 2, indicating that males either switch phenotypes with age or that Type 2 results in lower life expectancy, evidencing the costs of male reproduction in badgers.     

Individual differences in den maintenance effort in a communally dwelling mammal: the Eurasian badger

Group living has potential costs in terms of relative fitness for individuals that invest effort in activities providing general benefit, if other competing individuals exploit those activities and accrue similar benefits at no cost. We examined the roles of individual badgers, Meles meles, in the den maintenance activities of digging and bedding collection at their communal setts. Twenty per cent of adults and yearlings were responsible for 60-90% of the observed digging and bedding collection effort. Overall males tended to dig more than females, while durations of bedding collection were similar. Among adult and yearling males and females, individuals with a high percentage of days observed at the sett (high site fidelity) performed more digging and collected more bedding than transients and badgers of low site fidelity. Males of high status (large, mature, frequently copulating individuals) were more likely to dig than males of low status. Principal component analysis indicated negligibly low correlation between status and site fidelity for males. We hypothesize that while highly resident adult females benefit from extending the sett to avoid direct reproductive competition, males of high status and site fidelity might extend the sett to encourage receptive breeding females in their home group to stay and/or to improve survivorship of sired litters. Other categories of individual depend on the sett for shelter but, perhaps having less to gain from extending it, adopt a less active role in sett maintenance. Copyright 1999 The Association for the Study of Animal Behaviour.     

Increasing frequency of bite wounds with increasing population density in Eurasian badgers, Meles meles

Eurasian badgers sometimes live in territorial, mixed-sex groups; the adaptive significance of this is not understood, but members generally interact amicably. None the less, badgers occasionally fight and inflict sometimes severe wounds on one another. Based on 498 badger life histories, from first emergence as a cub until death, documented during a 10-year trapping study at Wytham Woods, Oxfordshire, U.K., the patterns and rates of bite wounding and consequential scarring were examined. Male badgers received more wounds and more severe wounds than did females. Wounding rates for both sexes increased significantly with age, and there was evidence that heavier individuals received most wounds. No seasonal pattern in wounding rates was apparent. During the study, the badger population size increased three-fold and wounding rates, particularly in males, showed a density-dependent increase. The rate of bite wounding increased with group size, and this increase was more marked among males than among females. Among males, but not females, the rate of bite wounding also increased with the number of badgers living in adjoining territories.     

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.     

Changes in the prevalence of badger persecution in Northern Ireland

Animal populations generally increase after release from hunting pressure and/or cessation of illegal persecution. Implementation of full legislative protection of the Eurasian badger Meles meles in Great Britain is thought to have led to increases in badger abundance due to reduced levels of persecution. Conversely, prevalence of badger persecution in Northern Ireland was historically much higher than in Great Britain, and badger abundance remained stable over time despite similar legislative protection. We examined temporal changes in the prevalence of badger sett disturbance in Northern Ireland from 1990/1993 to 2007/2008 in relation to population status. A total of 56 (12.6%) of 445 setts surveyed during 1990/1993 had been disturbed compared to 29 (4.4%) of 653 setts during 2007/2008. This was a significant decline (−65%) in the incidence of sett disturbance over the 14–18-year period. Most notably, the incidence of digging at badger setts, indicative of local badger baiting activity, declined from 50% to 3.5% of disturbed setts. Signs of recent disturbance were significantly more frequent at disused setts suggesting that once disturbed, badgers may vacate a sett. The number of badger social groups in Northern Ireland did not differ between the two study periods, suggesting that previously high levels of badger persecution did not limit the number of badger social groups. The stability of the badger population in Northern Ireland compared to the growing population in Great Britain cannot be attributed to changes in the prevalence of persecution. Differences in the trajectories of both populations could be due to a range of factors including climate, habitat composition and structure, farming practices or food availability. More work is needed to determine how such factors influence badger population dynamics.     

Culling-induced changes in badger (Meles meles) behaviour, social organisation and the epidemiology of bovine tuberculosis

In the UK, attempts since the 1970s to control the incidence of bovine tuberculosis (bTB) in cattle by culling a wildlife host, the European badger (Meles meles), have produced equivocal results. Culling-induced social perturbation of badger populations may lead to unexpected outcomes. We test predictions from the 'perturbation hypothesis', determining the impact of culling operations on badger populations, movement of surviving individuals and the influence on the epidemiology of bTB in badgers using data dervied from two study areas within the UK Government's Randomised Badger Culling Trial (RBCT). Culling operations did not remove all individuals from setts, with between 34-43% of badgers removed from targeted social groups. After culling, bTB prevalence increased in badger social groups neighbouring removals, particularly amongst cubs. Seventy individual adult badgers were fitted with radio-collars, yielding 8,311 locational fixes from both sites between November 2001 and December 2003. Home range areas of animals surviving within removed groups increased by 43.5% in response to culling. Overlap between summer ranges of individuals from Neighbouring social groups in the treatment population increased by 73.3% in response to culling. The movement rate of individuals between social groups was low, but increased after culling, in Removed and Neighbouring social groups. Increased bTB prevalence in Neighbouring groups was associated with badger movements both into and out of these groups, although none of the moving individuals themselves tested positive for bTB. Significant increases in both the frequency of individual badger movements between groups and the emergence of bTB were observed in response to culling. However, no direct evidence was found to link the two phenomena. We hypothesise that the social disruption caused by culling may not only increase direct contact and thus disease transmission between surviving badgers, but may also increase social stress within the surviving population, causing immunosuppression and enhancing the expression of disease.     

Serengeti real estate: density vs. fitness-based indicators of lion habitat quality

Habitat quality is typically inferred by assuming a direct relationship between consumer density and resource abundance, although it has been suggested that consumer fitness may be a more accurate measure of habitat quality. We examined density vs. fitness-based measures of habitat quality for lions in the Serengeti National Park, Tanzania. A 40-year average of female reproductive success (yearling cubs per female) was best explained by proximity to river confluences, whereas patterns of productivity (yearling cubs per km²) and adult female density (individuals per km²) were associated with more general measures of habitat quality and areas of shelter in poor habitat. This suggests that density may not accurately distinguish between high-quality 'source' areas and low-quality sites that merely provide refuges for effectively non-reproductive individuals. Our results indicate that density may be a misleading indicator of real estate value, particularly for populations that do not conform to an ideal free distribution.     

Assortative mating in fallow deer reduces the strength of sexual selection

Background

Assortative mating can help explain how genetic variation for male quality is maintained even in highly polygynous species. Here, we present a longitudinal study examining how female and male ages, as well as male social dominance, affect assortative mating in fallow deer (Dama dama) over 10 years. Assortative mating could help explain the substantial proportion of females that do not mate with prime-aged, high ranking males, despite very high mating skew. We investigated the temporal pattern of female and male matings, and the relationship between female age and the age and dominance of their mates.

Results

The peak of yearling female matings was four days later than the peak for older females. Younger females, and especially yearlings, mated with younger and lower-ranking males than older females. Similarly, young males and lower-ranking males mated with younger females than older males and higher-ranking males. Furthermore, the timing of matings by young males coincided with the peak of yearling female matings, whereas the timing of older male matings (irrespective of rank) coincided with the peak of older female matings.

Conclusions

Assortative mating, through a combination of indirect and/or direct female mate choice, can help explain the persistence of genetic variation for male traits associated with reproductive success.     

Annual and circadian activity patterns of badgers (Meles meles) in Białowieża Primeval Forest (eastern Poland) compared with other Palaearctic populations

Aim The annual and circadian rhythms and duration of activity of Eurasian badger Meles meles (Linnaeus 1758) were studied in a low‐density population inhabiting the primeval woodland in the European temperate zone. Results were compared with available data from the literature on seasonal changes in body mass and winter inactivity of badgers from across the Palaearctic region. Location Field work was carried out in Bia?owie?a Primeval Forest, eastern Poland. Biogeographical variation was reviewed based on twenty‐three localities in the Palaearctic region (from Western Europe to Central Siberia). Methods Thirteen badgers were radio‐collared in 1997–2001. Their circadian activity was sampled by 24‐h sessions of continuous radio‐tracking with location taken at 15‐min intervals. Annual activity was studied by radio‐tracking and inspections of setts. Earthworm (badgers’ main food) biomass was estimated in four types of habitats throughout the year. Results Badgers were nocturnal with one long bout of activity. Their rhythms of diel activity differed between spring and autumn, and between adult and subadult individuals. On average, badgers emerged from setts at 19:00 hours and returned to them at 03:42 hours. The highest level of activity was recorded between 20:00 and 03:00 hours. Duration of daily activity was, on average, 8.2 h day^?1, but varied significantly between seasons. The seasonal changes were inversely related to the abundance of earthworms. Duration of activity also depended on daily temperature, especially in the cold season. In winter, badgers stayed inactive for an average of 96 days per year. In autumn, they built fat reserves and their body mass nearly doubled compared with the spring values. The literature review on annual cycle of activity and body mass changes in Eurasian badgers showed that fat storage and duration of winter sleep strongly depended on climate (best approximated by January mean temperature). In regions with warm climates, badgers were active year round and their body mass changed only slightly, while in regions with severe winters badgers increased their body mass twofold from spring to autumn, and stayed inactive for as long as 6 months per year. Main conclusion We propose that, in the temperate and boreal zones of the Palaearctic region, the ultimate determinant of biogeographical variation in badgers’ annual activity is the winter shortage of earthworms, which are the main component of badger diet.     

Reproductive biology and ecology of female polar bears (Ursus maritimus)

Data on age-specific natality rates, litter size, interbirth interval, age of first reproduction, reproductive senescence, age of weaning and cub survival were determined for a free-ranging population of polar bears inhabiting Hudson Bay, Canada, near the southern limit of the species range. Serum progesterone levels were also determined for females at different stages of their reproductive cycle to provide corroborative support for the reproductive parameters described. Animals were live captured using immobilizing drugs and each animal uniquely marked for future identification. First parturition occurred at four or five years of age and the age-specific natality rate increased with age until approximately 20 years, after which it dropped markedly. At least 40% of adult females displayed two-year interbirth intervals and 55% of cubs in their second year were independent of their mother. Mean size of cub litters in spring was 1.9 and 13% of litters had three or more cubs. The natality rate for 5–20-year-old females was estimated as 0.9, higher than that reported for any more northerly polar bear populations where two-year interbirth intervals are rare, fewer than 5% of yearling cubs are weaned and triplet litters occur with less than 1% frequency. Cub mortality was initially high and declined with age. Although cubs in western Hudson Bay were weaned at a younger age and a lighter weight than their counterparts in more northern populations, cub mortality rates were similar. The reason for the marked differences in reproductive parameters in the western Hudson Bay population is not known. We speculate that sea-ice conditions may be sufficiently different to allow weaned bears at a lighter body weight to hunt seals more successfully there than further north.     

The distribution of Eurasian badger, Meles meles, setts in a high-density area: field observations contradict the sett dispersion hypothesis

Are setts significant determinants of badger socio‐spatial organisation, and do suitable sett sites represent a limited resource, potentially affecting badger distributions? The factors determining diurnal resting den, or sett, location and selection by Eurasian badgers Meles meles L. were investigated in Wytham Woods, Oxfordshire. 279 sett sites were located. The habitat parameters that were associated with the siting of these setts were analysed and associations were sought between sett location and character and the body condition and body weight of resident badgers Habitat characteristics in the vicinity of setts were significantly different from randomly selected points. Badgers preferentially selected sites with sandy, well‐drained soils, situated on NW‐facing, convex and moderately inclined slopes at moderate altitude. There was no evidence that sett morphology (number of entrances, sett area, number of hinterland latrines) was affected by the surrounding sett site habitat characteristics. Mean body weight was significantly higher for badgers occupying territories with setts in sandy soils, situated on NW‐facing slopes, than in territories with less optimal sett characteristics. Contrary to the hypothesis that the availability of sett sites was limiting, and therefore that sett dispersion dictates the spatial and social organisation of their populations, the badgers were clearly able to excavate new setts. On our measures, these new setts were not inferior to old established ones, despite occupying subsequently exploited sites; the badgers utilising these new setts had neither lighter body weights nor poorer body condition scores. During the period of our study badgers have manifestly been able to dig numerous new setts; as satisfactory sites still remain available, this indicates that suitable sett sites have not yet become a limiting resource. There was no relationship between sett age and the characteristics of the site in which it was dug, as suitable sites were not limiting. Significantly, population expansion during the decade 1987–1997 was not constrained by lack of setts, rather the main proliferation in setts occurred after the population size had peaked in 1996. Some implications for the management and conservation of the Eurasian badger are considered.