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.     

Comparative study on the consequences of culling badgers (Meles meles) on biometrics, population dynamics and movement

1.  Capture–mark–recapture data were used to describe the process of recovery from a typical badger removal operation (BRO) at North Nibley, Gloucestershire, UK, which was carried out as part of the government's strategy to control bovine tuberculosis. Data on biometrics, demographics and movement from this low-density disturbed population were compared with those of two nearby high-density undisturbed populations (Wytham Woods and Woodchester Park, UK) in order to study fundamental principles of population dynamics and density-dependence.
2.  Badgers moved more between social groups at North Nibley than in the other study areas, particularly in the immediate aftermath of the removal operation.
3.  Recolonization of the vacated habitat occurred in the first instance by young females.
4.  Although in the first year after the BRO no cubs had been reared in any of the culled groups, and although the shortage of sexually mature boars may have limited the reproductive output of sows in the following year, the population took only 3 years to recover to its (already lowered) preremoval density.
5.  Losses from the adult (and cub) population due to mortality or emigration were smaller at North Nibley than at the other sites.
6.  There was much evidence that during 1995 and 1996 density-dependent effects constrained the reproductive output of the high-density populations, and some support for the hypothesis that badgers exhibit the non-linear 'large mammal' type of functional response to density.
7.  Badgers at North Nibley were younger, heavier and in better condition than badgers at Wytham Woods and Woodchester Park.
8.  We argue that the disease dynamics are likely to be different in disturbed compared with undisturbed badger populations, and that this could affect the effectiveness of BROs.     

(Not) far from home: No sex bias in dispersal,but limited genetic patch size,in an endangered species,the Spotted Turtle (Clemmys guttata)

Sex-biased dispersal is common in many animals, with male-biased dispersal often found in studies of mammals and reptiles, including interpretations of spatial genetic structure, ostensibly as a result of male–male competition and a lack of male parental care. Few studies of sex-biased dispersal have been conducted in turtles, but a handful of studies, in saltwater turtles and in terrestrial turtles, have detected male-biased dispersal as expected. We tested for sex-biased dispersal in the endangered freshwater turtle, the spotted turtle (Clemmys guttata) by investigating fine-scale genetic spatial structure of males and females. We found significant spatial genetic structure in both sexes, but the patterns mimicked each other. Both males and females typically had higher than expected relatedness at distances <25 km, and in many distance classes greater than 25 km, less than expected relatedness. Similar patterns were apparent whether we used only loci in Hardy–Weinberg equilibrium (n = 7) or also included loci with potential null alleles (n = 5). We conclude that, contrary to expectations, sex-biased dispersal is not occurring in this species, possibly related to the reverse sexual dimorphism in this species, with females having brighter colors. We did, however, detect significant spatial genetic structure in males and females, separate and combined, showing philopatry within a genetic patch size of <25 km in C. guttata, which is concerning for an endangered species whose populations are often separated by distances greater than the genetic patch size.     

Genetic structure of Eurasian badgers Meles meles (Carnivora: Mustelidae) and the colonization history of Ireland

The present study examined the contemporary genetic composition of the Eurasian badger, Meles meles, in Ireland, Britain and Western Europe, using six nuclear microsatellite loci and a 215‐bp fragment of the mitochondrial DNA control region. Significant population structure was evident within Europe (global multilocus microsatellite F_ST = 0.205, P < 0.001; global mitochondrial control region Φ_ST = 0.399, P < 0.001). Microsatellite‐based cluster analyses detected one population in Ireland, whereas badgers from Britain could be subdivided into several populations. Excluding the island populations of Ireland and Britain, badgers from Western Europe showed further structuring, with evidence of discrete Scandinavian, Central European, and Spanish populations. Mitochondrial DNA cluster analysis grouped the Irish population with Scandinavia and Spain, whereas the majority of British haplotypes grouped with those from Central Europe. The findings of the present study suggest that British and Irish badger populations colonized from different refugial areas, or that there were different waves of colonization from the source population. There are indications for the presence of an Atlantic fringe element, which has been seen in other Irish species. We discuss the results in light of the controversy about natural versus human‐mediated introductions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Morphological variability and developmental instability in subpopulations of the Eurasian badger (Meles meles) in Denmark

Aim Local populations from different geographical regions may differ in the selection regimes to which they are exposed. Differences in environmental factors and population density may affect the relative importance of different selective forces (e.g. natural vs. sexual selection). We suggest a direction of investigation concerned with the developmental instability of morphological traits. The goal is to disclose putative small‐scale geographical differences in the evolutionary forces, which may be hard to detect. Location Craniometrical investigations were carried out on ninety‐eight skulls and teeth of the Eurasian badger (Meles meles) collected during the period 1995–97 from three different populations in Denmark. One of these thrives at low population density, whereas the two others are characterized by high local density. Methods The skulls were investigated for developmental instability (DI) using fluctuating asymmetry (FA) as its estimator. FA was measured on canines, molars, premolar teeth and other skull and mandible traits. For the statistical analyses, we applied nonparametric permutation tests. Results Evidence was found suggesting differentiation among populations in mean degree of FA, and the FA values measured on canines were higher in the high‐density populations. FA of the canines was significantly higher in males than females, in contrast to FA of the other traits. Evidence of a negative relationship between canine size and their FA was found, whereas no significant correlations were found between the molar and premolar teeth measures and their FA. Main conclusions Our results suggest that canines could be under directional selection stemming from intrasexual competition, which may be stronger in high‐density zones. The other teeth investigated seem to be under a stabilizing regime hence their FA is mainly affected by environmental stresses. The negative relationship between canine size and FA found in males suggests the capacity of badgers to respond in an evolutionary way to environmental changes, despite the low genetic variability previously found at the molecular level.     

Social organization and movement influence the incidence of bovine tuberculosis in an undisturbed high-density badger Meles meles population

1. The culling of European badgers Meles meles has been a central part of attempts to control bovine tuberculosis (TB) in British cattle for many years. Recent results, however, indicate that this approach could in practice enhance disease spread. 2. This paper looks at the relationship between TB incidence and badger ecology in a high-density population in south-west England, which has been the subject of a long-term intensive study. The principal aims were to relate the probability of TB incidence, as detected by culture of clinical samples (i.e. excretion of bacilli), at the level of the individual and of the social group to demographic processes, movement, social organization and disease dynamics. 3. The probability of an individual being an incident case was greater in groups where TB was already present, although this was less influential in groups that were subject to some instability in numbers. Both individuals and groups were more likely to be incident cases where the social group was diminishing in size, although no relationship was observed with group size itself. This suggests that the process of group size reduction rather than group size per se has most influence on disease dynamics. The likelihood that either an individual or a group was an incident case was positively correlated with both individual and group-level movement. When the proportion of females in a social group was high, the positive association between movement and incidence was found to be more pronounced and there was a significantly higher probability of incident cases among males. 4. These relationships highlight the importance of social structure in driving TB transmission dynamics in this stable, high-density badger population. The results support the idea that a stable social structure mitigates against new incident cases of disease, and are consistent with the contention that badger culling may create the social circumstances for enhanced transmission of TB.     

Isolation by distance and gene flow in the Eurasian badger (Meles meles) at both a local and broad scale

Eurasian badgers, Meles meles, have been shown to possess limited genetic population structure within Europe; however, field studies have detected high levels of philopatry, which are expected to increase population structure. Population structure will be a consequence of both contemporary dispersal and historical processes, each of which is expected to be evident at a different scale. Therefore, to gain a greater understanding of gene flow in the badger, we examined microsatellite diversity both among and within badger populations, focusing on populations from the British Isles and western Europe. We found that while populations differed in their allelic diversity, the British Isles displayed a similar degree of diversity to the rest of western Europe. The lower genetic diversity occurring in Ireland, Norway and Scotland was more likely to have resulted from founder effects rather than contemporary population density. While there was significant population structure (F ST = 0.19), divergence among populations was generally well explained by geographic distance (P < 0.0001) across the entire range studied of more than 3000 km. Transient effects from the Pleistocene appear to have been replaced by a strong pattern of genetic isolation by distance across western Europe, suggestive of colonization from a single refugium. Analysis of individuals within British populations through Mantel tests and spatial autocorrelation demonstrated that there was significant local population structure across 3-30 km, confirming that dispersal is indeed restricted. The isolation by distance observed among badger populations across western Europe is likely to be a consequence of this restricted local dispersal.     

Population genetic structure, gene flow and sex-biased dispersal in frillneck lizards (Chlamydosaurus kingii)

By using both mitochondrial and nuclear multiloci markers, we explored population genetic structure, gene flow and sex-specific dispersal of frillneck lizards ( Chlamydosaurus kingii ) sampled at three locations, separated by 10 to 50 km, in a homogenous savannah woodland in tropical Australia. Apart from a recombinant lizard, the mitochondrial analyses revealed two nonoverlapping haplotypes/populations, while the nuclear markers showed that the frillneck lizards represented three separate clusters/populations. Due to the small population size of the mtDNA, fixation may occur via founder effects and/or drift. We therefore suggest that either of these two processes, or a combination of the two, are the most likely causes of the discordant results obtained from the mitochondrial and the nuclear markers. In contrast to the nonoverlapping mitochondrial haplotypes, in 12 out of 74 lizards, mixed nuclear genotypes were observed, hence revealing a limited nuclear gene flow. Although gene flow should ultimately result in a blending of the populations, we propose that the distinct nuclear population structure is maintained by frequent fires resulting in local bottlenecks, and concomitant spatial separation of the frillneck lizard populations. Limited mark–recapture data and the difference in distribution of the mitochondrial and nuclear markers suggest that the mixed nuclear genotypes were caused by juvenile male-biased dispersal.     

Spatial population genetic structure in Trillium grandiflorum: the roles of dispersal, mating, history, and selection

The roles of the various potential ecological and evolutionary causes of spatial population genetic structure (SPGS) cannot in general be inferred from the extant structure alone. However, a stage-specific analysis can provide clues as to the causes of SPGS. We conducted a stage-specific SPGS analysis of a mapped population of about 2000 Trillium grandiflorum (Liliaceae), a long-lived perennial herb. We compared SPGS for juvenile (J), nonreproductive (NR), and reproductive (R) stages. Fisher's exact test showed that genotypes had Hardy-Weinberg frequencies at all loci and stage classes. Allele frequencies did not differ between stages. Bootstrapped 99% confidence intervals (99%CI) indicate that F-statistic values are indistinguishable from zero, (except for a slightly negative FIT for the R stage). Spatial autocorrelation was used to calculate f the average kinship coefficient between individuals within distance intervals. Null hypothesis 99%CIs for f were constructed by repeatedly randomizing genotypic locations. Significant positive fine-scale genetic structure was detected in the R and NR stages, but not in the J stage. This structure was most pronounced in the R stage, and declined by about half in each remaining stage: near-neighbor f = 0.122, 0.065, 0.027, for R, NR, and J, respectively. For R and NR, the near-neighbor f lies outside the null hypothesis 99%CI, indicating kinship at approximately the level of half-sibs and first cousins, respectively. We also simulated the expected SPGS of juveniles post dispersal, based on measured R-stage SPGS, the mating system, and measured pollen and seed dispersal properties. This provides a null hypothesis expectation (as a 99%CI) for the J-stage correlogram, against which to test the likelihood that post-dispersal events have influenced J-stage SPGS. The actual J correlogram lies within the null hypothesis 99%CI for the shortest distance interval and nearly all other distance intervals indicating that the observed low recruitment, random mating and seed dispersal patterns are sufficient to account for the disappearance of SPSG between the R and the J stages. The observed increase in SPGS between J and R stages has two potential explanations: history and local selection. The observed low total allelic diversity is consistent with a past bottleneck: a possible historical explanation. Only a longitudinal stage-specific study of SPGS structure can distinguish between historical events and local selection as causes of increased structure with increasing life history stage.     

Net costs of group living in a solitary forager, the Eurasian badger (Meles meles)

Group living is expected to evolve under two extreme conditions:when there are net benefits to the individual of living in agroup and when there are net costs to living in a group togetherwith strong ecological constraints on dispersal and independentbreeding. A third condition may facilitate group formation:when the territorial defense of resources that are dispersedwidely or renewed at high rates allows groups to form at nocost to group members. Eurasian badgers are solitary over alarge part of their geographic distribution and forage aloneeven where diey live in territorial groups. To determine whichof the three conditions best explains group living in badgers,we analyzed dispersal and breeding patterns. Using these datawe describe the mechanism of group formation and the net costsor benefits of group living in 16 badger groups in one population.Groups form primarily by the retention of young on their natalterritory. Typically, groups contain one or more sexually maturebut nonbreeding females in addition to one or more breedingfemales. Females do not benefit from group living as indicatedby the lack of relationships, or perhaps negative relationships,between the proportion of adult females that bred in a groupand mean reproductive success, on the one hand, and group sizeand components of group size, on the other. A high reproductivefailure rate among females in this and other populations ofsocial badgers, as compared with a population of solitary badgers,suggests that there is a cost of group living to females. Wepropose that if badgers defend resources whose spatial distributionor high renewal rates allow groups to form on a territory atlitde or no cost to group members, weak ecological constraintson dispersal and independent breeding are sufficient to explainthe formation of groups. Badgers appear to have not evolvedcooperative breeding in response to these ecological conditions,perhaps because badger groups represent an early stage in theevolution of carnivore sociality.     

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.     

Evidence that disease-induced population decline changes genetic structure and alters dispersal patterns in the Tasmanian devil

Infectious disease has been shown to be a major cause of population declines in wild animals. However, there remains little empirical evidence on the genetic consequences of disease-mediated population declines, or how such perturbations might affect demographic processes such as dispersal. Devil facial tumour disease (DFTD) has resulted in the rapid decline of the Tasmanian devil, Sarcophilus harrisii, and threatens to cause extinction. Using 10 microsatellite DNA markers, we compared genetic diversity and structure before and after DFTD outbreaks in three Tasmanian devil populations to assess the genetic consequences of disease-induced population decline. We also used both genetic and demographic data to investigate dispersal patterns in Tasmanian devils along the east coast of Tasmania. We observed a significant increase in inbreeding (F_IS pre/post-disease −0.030/0.012, P<0.05; relatedness pre/post-disease 0.011/0.038, P=0.06) in devil populations after just 2–3 generations of disease arrival, but no detectable change in genetic diversity. Furthermore, although there was no subdivision apparent among pre-disease populations (θ=0.005, 95% confidence interval (CI) −0.003 to 0.017), we found significant genetic differentiation among populations post-disease (θ=0.020, 0.010–0.027), apparently driven by a combination of selection and altered dispersal patterns of females in disease-affected populations. We also show that dispersal is male-biased in devils and that dispersal distances follow a typical leptokurtic distribution. Our results show that disease can result in genetic and demographic changes in host populations over few generations and short time scales. Ongoing management of Tasmanian devils must now attempt to maintain genetic variability in this species through actions designed to reverse the detrimental effects of inbreeding and subdivision in disease-affected populations.     

Estimating seed vs. pollen dispersal from spatial genetic structure in the common ash

Spatial genetic structure was analysed with five highly polymorphic microsatellite loci in a Romanian population of common ash (Fraxinus excelsior L.), a wind-pollinated and wind-dispersed tree species occurring in mixed deciduous forests over almost all of Europe. Contributions of seed and pollen dispersal to total gene flow were investigated by analysing the pattern of decrease in kinship coefficients among pairs of individuals with geographical distance and comparing it with simulation results. Plots of kinship against the logarithm of distance were decomposed into a slope and a shape component. Simulations showed that the slope is informative about the global level of gene flow, in agreement with theoretical expectations, whereas the shape component was correlated with the relative importance of seed vs. pollen dispersal. Hence, our results indicate that insights into the relative contributions of seed and pollen dispersal to overall gene flow can be gained from details of the pattern of spatial genetic structure at biparentally inherited loci. In common ash, the slope provided an estimate of total gene dispersal in terms of Wright's neighbourhood size of Nb = 519 individuals. No precise estimate of seed vs. pollen flow could be obtained from the shape because of the stochasticity inherent to the data, but the parameter combinations that best fitted the data indicated restricted seed flow, sigmas pound 14 m, and moderate pollen flow, 70 m pound sigmap pound 140 m.     

Determining natal dispersal patterns in a population of North American pikas (Ochotona princeps) using direct mark-resight and indirect genetic methods

A combination of direct (mark-resight) and indirect geneticmethods were used to investigate natal dispersal patterns andgenetic population structure in a population of North Americanpikas, Ochotona princeps. Pikas are small lagomorphs found intalus habitat of alpine areas throughout western North America.Adult pikas are individually territorial and rarely disperse.I used multilocus DNA fingerprinting to identify the parentsof juvenile animals. The settlement pattern of marked juvenilesand the pattern of relatedness of pikas across the study sitewas then examined within the study area. Although juvenilesborn at the study site exhibited a philopatric settlement pattern,an isolation-by-distance analysis did not reveal clusters ofhighly related individuals within the population. The F_ST estimatesuggests little genetic differentiation between populations2 km apart, and average DNA fingerprinting band-sharing amongadults was similar to values reported for outbred species. Anaverage of 34% of the adult population was replaced each winterby immigrants. DNA fingerprinting band-sharing analysis suggeststhat these immigrants had dispersed short, intermediate, andlong distances. These findings differ from earlier studies whichused observations of marked animals only to characterize dispersalpatterns. Direct observations of marked juveniles had documenteda philopatric settlement pattern, little or no dispersal outof natal populations, and no direct evidence of long distancemovement. Of the three major hypotheses proposed to explainthe evolution of dispersal in birds and mammals, competitionfor resources, competition for mates, and inbreeding avoidance,the results of this study support a competition for resourceshypothesis, where the key resource is territory     

Effects of population characteristics and structure on estimates of effective population size in a house sparrow metapopulation

Effective population size (N_e) is a key parameter to understand evolutionary processes and the viability of endangered populations as it determines the rate of genetic drift and inbreeding. Low N_e can lead to inbreeding depression and reduced population adaptability. In this study, we estimated contemporary N_e using genetic estimators (LDNE, ONeSAMP, MLNE and CoNe) as well as a demographic estimator in a natural insular house sparrow metapopulation. We investigated whether population characteristics (population size, sex ratio, immigration rate, variance in population size and population growth rate) explained variation within and among populations in the ratio of effective to census population size (N_e/N_c). In general, N_e/N_c ratios increased with immigration rates. Genetic N_e was much larger than demographic N_e, probably due to a greater effect of immigration on genetic than demographic processes in local populations. Moreover, although estimates of genetic N_e seemed to track N_c quite well, the genetic N_e‐estimates were often larger than N_c within populations. Estimates of genetic N_e for the metapopulation were however within the expected range (<N_c). Our results suggest that in fragmented populations, even low levels of gene flow may have important consequences for the interpretation of genetic estimates of N_e. Consequently, further studies are needed to understand how N_e estimated in local populations or the total metapopulation relates to actual rates of genetic drift and inbreeding.     

Spatial genetic structure in Milicia excelsa (Moraceae) indicates extensive gene dispersal in a low-density wind-pollinated tropical tree

In this study, we analysed spatial genetic structure (SGS) patterns and estimated dispersal distances in Milicia excelsa (Welw.) C.C. Berg (Moraceae), a threatened wind-pollinated dioecious African tree, with typically low density (∼10 adults/km²). Eight microsatellite markers were used to type 287 individuals in four Cameroonian populations characterized by different habitats and tree densities. Differentiation among populations was very low. Two populations in more open habitat did not display any correlation between genetic relatedness and spatial distance between individuals, whereas significant SGS was detected in two populations situated under continuous forest cover. SGS was weak with a maximum S _p-statistic of 0.006, a value in the lower quartile of SGS estimates for trees in the literature. Using a stepwise approach with Bayesian clustering methods, we demonstrated that SGS resulted from isolation by distance and not colonization by different gene pools. Indirect estimates of gene dispersal distances ranged from σ _g = 1 to 7.1 km, one order of magnitude higher than most estimates found in the literature for tropical tree species. This result can largely be explained by life-history traits of the species. Milicia excelsa exhibits a potentially wide-ranging wind-mediated pollen dispersal mechanism as well as very efficient seed dispersal mediated by large frugivorous bats. Estimations of gene flow suggested no major risk of inbreeding because of reduction in population density by exploitation. Different strategy of seed collection may be required for reforestation programmes among populations with different extent of SGS.