A survey of helminth infection in Eurasian badgers (Meles meles) in relation to their foraging behaviour in a Mediterranean environment in southwest Portugal

This study provides the first data on the helminth fauna of the Eurasian badger in the southwestern edge of its range (Grândola Mountain, Portugal) and interprets the results in relation to badger diet and feeding behaviour. By examination of 163 badger faecal samples, faecal developmental stages (eliminative forms) of four helminth species and one genus were identified: one cestode (Atriotaenia incisa) and four nematodes (Mastophorus muris, Molineus patens, Uncinaria criniformis and Strongyloides sp.). The overall prevalence of parasites was 62%, with limited seasonal variation. Single parasite excretions were dominant and Strongyloides sp. excretion was the most common. Diet assessment based on 450 faecal samples revealed that badgers consumed mainly insects and fruits. No correlation was detected between helminth prevalence and diet. Apparently, diet (mainly insects) and feeding behaviour (fossorial), together with the species’ social behaviour (anal scent marking of group members), facilitate the infection with helminths. The helminth fauna of Eurasian badgers in Grândola Mountain has isolationist characteristics, apparently indicating low host colonisation.     

Scent-marking behaviour of the honey badger, Mellivora capensis (Mustelidae), in the southern Kalahari

We investigated sexual and seasonal patterns in scent-marking behaviour of the honey badger, by direct observations of habituated individuals (five females, four adult males, two young males). Four categories of scent-marking behaviour were identified: (1) scent marking at latrines; (2) token urination in holes along the foraging path; (3) squat marking at single-use sites; and (4) functional excretion. Females and young males used all four types of scent marking, but adult males were not observed to use token urination. A strategy of hinterland scent marking was used, as was predicted from the large home ranges of both male and female honey badgers. There were significant sexual differences in marking rate: adult males primarily used latrines and adult females favoured token urination. Latrine scent marking in adult male honey badgers provides support for the ‘scent-matching’ hypothesis. Females visited latrines when they were in oestrus. However, the low level of marking activity during a visit and the intensive smelling suggested a scent-matching function rather than reproductive advertisement. Token urination appeared to be related to the maintenance of spatiotemporal separation in females, although we also observed token urination in young males. While the placement of urine in foraging holes and its relation with successful digging attempts offer some support for the foraging efficiency hypothesis, we consider this unlikely, because we did not observe it in adult males and there was no seasonal pattern. Squat marking occurred under a wide range of conditions in both males and females and may be related to marking valuable resources. It is likely that scent marking in honey badgers has many functions.     

Defecation and urination patterns of badgersMeles meles at low density in south west England

The spatial distribution of urine and faecal scent marks of badgersMeles meles (Linnaeus, 1758) at low population density (mean±SE across 4 social groups was 5.73±0.735 badgers/km2) in south-western England were quantified. Eighteen badger latrines (greater than one dung pit containing faeces), 74 single defecations not in pits and 21 faeces in single pits were located in spring when badgers were defending well-defined territories. Woodland was selected, and arable land avoided, for latrine sites. Pasture and built-up land was selected for single defecations not in pits whereas faeces in single pits were distributed randomly across habitat types. Faecal scent marks were strongly associated with the edge of pastoral fields rather than the middle. Forty-six and 51 urinations were located in spring and summer, respectively. Urine was deposited randomly across habitat types but was concentrated at the linear features surrounding the main setts. This is the first reported use of high levels of single defecations and urinations in badger scent marking strategies in the UK. These results are discussed in relation to the potential for transmission of bovine tuberculosisMycobacterium bovis from badger excreta to cattle.     

Lunar-related reproductive behaviour in the badger (Meles meles)

Cyclicity in behaviours, including reproduction, in relation to the lunar cycle is widely documented in some phyla, but weak or unknown in Class Mammalia. In this paper we present long-term video surveillance data of wild Eurasian badgers Meles meles, which reveal a strong correlation between reproductive behaviour and the lunar cycle. Squat marking and raised-leg urination, which increase in frequency at times of reproductive activity, showed maxima around day 22 of the lunar cycle (i.e. new moon). These findings were supported by observations of matings, together with published records, which showed significantly higher occurrence in the lunar dark phase (last quarter to first quarter). We propose that the lunar cycle has the potential to act as a regulator of the reproductive cycle in the badger.     

The behavioural responses of badgers (Meles meles) to exclusion from farm buildings using an electric fence

Behavioural investigations into the transmission of bovine tuberculosis (Mycobacterium bovis) between badgers and cattle suggest that badger activity in farm buildings may incur a significant risk of cross-infection. However, measures to exclude badgers from buildings have not been systematically field-tested. In the present study, remote surveillance and radio-tracking were used to monitor the effect of electric fencing manipulations on the frequency of badger incursions into feed stores and cattle housing, and on badger ranging behaviour. Electric fencing was effective in preventing access to the farm buildings where it was installed and also significantly reduced incursions into unfenced buildings. Badger home range and core activity areas tended to increase in size when the fencing was installed, although they did not extend beyond the boundaries of the relevant social group territories. We discuss the logistical constraints of using electric fencing in this context and conclude that it is a potentially useful method of reducing contact between badgers and cattle, within farm buildings and yards.     

How many Eurasian badgers <Emphasis Type="Italic">Meles meles</Emphasis> L. are there in the Republic of Ireland?

In Ireland, the badger Meles meles L is a reservoir species for Mycobacterium bovis and, as such, contributes to the maintenance of bovine tuberculosis in cattle. A previous estimate of the badger population in the Republic was 200,000 badgers. In the current study, we obtained data on badger numbers from a large-scale badger removal project (the Four-Area project). The removal areas of the Four-Area Project were surrounded by barriers (either water or buffer areas where removals were also conducted) to prevent badger immigration. Within these areas, a grid of 0.25 km² was created within which we knew the badger numbers and habitat types (based on Corine data). Associations between badger numbers and habitat type were investigated using negative binomial modeling. Extrapolations from the model yielded an estimated badger population in the Republic of approximately 84,000 badgers. The implications of these findings are discussed.     

The behavioural response of the white-clawed crayfish (Austropotamobius pallipes) to five standard marking techniques over a 14-day period

ABSTRACT

The white-clawed crayfish (Austropotamobius pallipes), is classified as endangered in the 2010 IUCN Red List and is protected under Irish and European legislation due to a drastic decline in abundance and range over the past century. Mark-recapture methods play a vital role in the process of estimating population size, and in the monitoring and conservation of mobile species. A variety of marking techniques are routinely used in a range of crayfish species for ecological research. The majority of information on the effects of these marking techniques on the physiology of crayfish species focuses on survivability and growth, whereas information is scarce on the effects on crayfish behaviour. This study investigates the behavioural changes of crayfish in reaction to five commonly used marking techniques in laboratory conditions. The marking techniques used were internal and external passive integrated transponder (PIT) tags, visible implant elastomer (VIE) tags, cauterisation and ablation. Effects of marking technique on behaviour were evaluated immediately after marking, and over a 14-day period, on 78 crayfish. Results indicated that the internal passive integrated transponder (PIT) tags negatively affected crayfish behaviour by increasing resting levels. Ablation and cauterisation also increased grooming levels, while marking reduced activity levels in general. These effects on behaviour occurred immediately after marking, but quickly dissipated. These results suggest that the use of internal PIT tags for marking white-clawed crayfish may need to be avoided as increased resting levels could lead to increased exposure out of refuge and predation. This may have a detrimental impact on the welfare and conservation of white-clawed crayfish and also lead to biased population estimates during mark-recapture projects. The remaining marking techniques (external PIT tag, VIE tag, ablation, cauterisation) may be used, as changes in behaviour will not have a detrimental impact and therefore should not lead to biased population estimates.     

Distribution of badger latrines in a high-density population: habitat selection and implications for the transmission of bovine tuberculosis to cattle

Eurasian badgers Meles meles habitually deposit droppings and other scent marks at latrines, which may be associated with territorial defence, and communicate information related to group and individual identity and status, and food resources. Understanding patterns of latrine distribution contributes to our understanding of badger social behaviour, and may be relevant to managing the risks of transmission of bovine tuberculosis from badgers to cattle. We investigated the distribution of badger latrines relative to habitat composition in a high-density badger population occupying a 7 km² area of diverse landscape in south-west England. Results indicated that the frequency and density of badger latrines varied according to land use, with woodland and linear landscape features (particularly hedges and stone walls) being positively selected. The number of latrines decreased significantly with distance from linear features. Grassland was negatively selected given its availability, but contained the highest number of latrines. The tendency for latrines to be associated with particular habitat types covaried spatially across the study area. We present a habitat selection probability function, based on the output of our analyses, to allow comparison of observed versus expected latrine counts per habitat type at different sites. Habitat manipulation on farmland may offer opportunities to manage exposure of cattle to badger latrines. However, our analyses indicate that other factors (perhaps demographic or environmental) may also exert a substantial local influence on latrine location.     

Badgers and Badger Fleas: Strategies and Counter-Strategies

We examined the broad hypothesis that one function of grooming by the European badger (Meles meles) is to disadvantage (possibly by removal) parasitic badger fleas (Paraceris melis). We pursued two lines of investigation. First, we used infra‐red video analysis to examine the body areas reached by self‐ and allo‐grooming badgers. We expected that if grooming was important to disadvantage fleas then allo‐grooming would cover areas that could not be reached by self‐grooming. Badgers preferred dorsal allo‐grooming and ventral self‐grooming, and when combined, the overall amount of grooming per square centimetre of skin area provided even body coverage, pointing to a hygienic function, rather than a purely social function. Secondly, we examined fleas’ responses to simulated fur disturbance, characteristic of grooming. If grooming had no flea disadvantage effect, we would expect no response from fleas, and no directionality in their movement away from direct touch. The number of fleas encountered rapidly declined in successive 10‐s counts during simulated ‘grooming’ at the same site. When badgers were ‘groomed’ on alternate sides (mimicking the badgers’ rapid alternation of grooming position), there was a marked increase in fleas when grooming resumed on the original side. Similarly, when ‘grooming’ was suspended for 40 s, there was an initial increase in the number of fleas when ‘grooming’ was resumed. Disturbed fleas tended to run downwards relative to gravity and towards the posterior of the badger, following the direction of hair growth. This contrasted with the behaviour of fleas removed from badgers which tended to run upwards and jump. We concluded that the pattern of badger grooming and the fleas’ response to disturbance was consistent with a hypothesis that badgers and badger fleas have strategies and counter strategies to maximize and minimize contact (respectively) during grooming.     

Interspecific variation of scent-marking behaviour in wild tamarins, Saguinus mystax and Saguinus fuscicollis.

The scent-marking behaviour of sympatric moustached, Saguinus mystax, and saddle-back tamarins, Saguinus fuscicollis, was compared in order to explore interspecific differences and potential sources of variation. The author examined basic patterns of scent marking (types, intensity, complexity), substrate use (type, orientation, height), and social patterning of scent marking in three groups of S. mystax and one group of S. fuscicollis at the Estación Biológica Quebrada Blanco, Peruvian Amazonia. S. mystax and S. fuscicollis differed significantly in the relative frequency of different types, and in the intensity and complexity of scent marking. Only S. fuscicollis showed allomarking. They also differed significantly in the type, orientation and height of substrates used for scent marking which corresponded to general differences in substrate use. In S. fuscicollis, but very rarely in S. mystax, two or more group members marked the same site sequentially or simultaneously. 'Collective scent marking', i.e. simultaneous scent marking by most or all group members, occurred only in S. fuscicollis. Since both tamarin species live sympatrically in mixed-species groups, ecological factors are unlikely to account for the differences found in scent-marking behaviour (except for differences in substrate use). They probably relate to as yet unknown differences in social and reproductive strategies of the two species.     

The Equivocal Relationship Between Territoriality and Scent Marking in Wild Saddleback Tamarins (<Emphasis Type="Italic">Saguinus fuscicollis</Emphasis>)

Researchers have often assumed that scent marking serves a territorial function in callitrichines, although some controversy exists. To fulfill such a function, scent marks should 1) prevent intrusions, 2) ensure access to feeding resources, 3) enable avoidance of intergroup encounters, or 4) play an important role in the aggressive encounters between groups. We studied 13 saddleback tamarins (Saguinus fuscicollis) belonging to 3 free-ranging groups, which formed mixed-species troops with moustached tamarins (S. mystax) in the Amazonian rain forest of Peru. None of the predictions were confirmed. The tamarins used a border-marking strategy, marking more on the periphery of their territory. However, feeding trees in overlap and encounter areas received more scent marking but were still visited by neighboring groups. Intergroup encounters occurred more often than expected, and scent-marking frequency was not higher during them than when no other group was present. It appears that instead of defending a territory in the classic sense, the tamarins are optimizing signal transmission by depositing their scents where the probability of detection by neighbors is higher. Saddleback tamarins may use shared areas of their home ranges to exchange information with neighboring groups, perhaps regarding reproductive opportunities.     

Dominance hierarchy and space use pattern in male captive muntjacs,Muntiacus reevesi

Social behaviour and space use patterns of 15 male muntjacs (Muntiacus reevesi) were studied in a large enclosure in 1979 and 1980. Males often interacted aggressively with each other showing distinct aggressive behaviour patterns. A dominance hierarchy was established and 4 adult males emerged as co-dominant. Movements of these dominant males were confined within small areas which did not overlap with each other and partitioned the enclosure. They defended and maintained the areas by aggressive behaviour and scent marking.     

Marking as a decision factor in the choice of a new resting site in <Emphasis Type="Italic">Lasius niger</Emphasis>

Summary In ants, aggregation is mainly based on the attraction between individuals. We confirmed the existence of ground marking in Lasius niger and demonstrated its influence on the aggregative behaviour of ants: the place of the gathering is leading by the existence of a ground marking which favours clustering by increasing the rate of the process but does not change the aggregation level. This factor could play a role in the choice of a new nest site and in the spatial distribution of ants inside the nest. This marking, presuming to have a long duration, could represent an external memory which could play an important role in various situations such as the choice of new nest or the reorganisation of ants after a disturbance.Received 23 July 2003; revised 16 December 2003; accepted 19 December 2003.     

How does food distance influence foraging in the ant Lasius niger: the importance of home-range marking

We study the influence of food distance on the individual foraging behaviour of Lasius niger scouts and we investigate which cue they use to assess their distance from the nest and accordingly tune their recruiting behaviour. Globally, the number of U-turns made by scouts increases with distance resulting in longer travel times and duration of the foraging cycle. However, over familiar areas, home-range marking reduces the frequency and thereby the impact of U-turns on foraging times leading to a quicker exploitation of food sources than over unmarked set-ups. Regarding information transfer, the intensity of the recruitment trail reaching the nest decreases with increasing food distance for all set-ups and is even more reduced in the absence of home-range marking. Hence, the probability of a scout continuing to lay a trail changes along the homeward journey but in a different way according to home-range marking. Over unexplored setups, at a given distance from the food source, the percentage of returning trail-laying ants remains unchanged for all tested nest-feeder distances. Hence, the tuning of the trail recruiting signal by scouts was not influenced by an odometric estimate of the distance already travelled by the ants during their outward journey to the food. By contrast, over previously explored set-ups, a distance-related factor – that is the intensity of home-range marking – strongly influences their recruiting behaviour. In fact, over a home-range marked bridge, the probability of returning ants maintaining their trail-laying behaviour increases with decreasing food distance while the gradient of home-range marks even induces ants which have stopped laying a trail to resume this behaviour in the nest vicinity. We suggest that home-range marking laid passively by walking ants is a relevant cue for scouts to indirectly assess distance from the nest but also local activity level or foraging risks in order to adaptively tune trail recruitment and colony foraging dynamics. Received 13 July 2004; revised 26 January and 20 May 2005; accepted 2 July 2005.     

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.     

Variation in stimulus,seasonal context,and response to urine marks by captive Iberian wolves (<Emphasis Type="Italic">Canis lupus signatus</Emphasis>)

Urine marking behaviour was observed over 14 months in two captive pairs of Iberian wolves (Canis lupus signatus) by means of behaviour sampling. The study focused on the relative attractiveness of the different stimuli toward which urinations are directed, the seasonal context of urine marking, and the response by mates to sexually dimorphic action patterns. Kinds of marked substrates varied according to the posture used by wolves to urinate [raised-leg urination (RLU), flexed-leg urination (FLU), standing urination (STU), or squat urination (SQU)]. In RLUs and FLUs the wolves mainly used conspicuous substrates, whereas STUs and SQUs were carried out directly onto inconspicuous substrates. Wolves urinated on plants more than on other substrates, mostly selecting trees. A selection of trees according to their trunk diameter was also observed. The posts were chosen or avoided seemingly because of their specific characteristics such as size. In short, wolves marked with urine (RLUs and FLUs) those substrates that secured a greater effectiveness of marking. The mean duration of RLUs was lower than that of FLUs and SQUs. The rate of RLUs in males increased in winter (November–December) and in summer (July–August), while the rate of FLUs and SQUs in females and STUs in males did not increase during these periods. The wolves investigated RLUs more frequently than SQUs. The RLU display appears to be more ancestral than derived, similar in all species of canids and even in the Iberian wolf.     

Predicting Distribution and Density of European Badger (<Emphasis Type="Italic">Meles Meles</Emphasis>) Setts in Denmark

Predictive models of the spatial distribution and abundance of species based on habitat characteristics are finding increasing use in management and conservation. The European badger attracts interest as a model species both for conservation reasons and because of the important role the species is playing in understanding carnivore sociality. We developed a statistical habitat model based on presence/absence data on badger setts. To maximise the utility of the model in management, we limited the choice of model variables to those that had a clear basis in badger ecology and that could be obtained on a nation-wide digital format. We extrapolated the habitat model to a region in Denmark and developed a threshold-independent sett distribution algorithm to estimate sett densities. The habitat model was simpler than previously published models of badger sett habitat selection, but nevertheless had a predictive ability in excess of 80% judged against independent data. The sett distribution algorithm was able to simultaneously reproduce several observed patterns of sett density and distribution over the probability gradient. It thus represents a significant improvement over threshold-dependent methods used to discriminate between suitable and unsuitable habitat predicted by presence/absence regression models. Our approach demonstrates that a model of badger sett habitat suitability with high predictive power can be obtained using easily accessible map-variables and presence/absence data. This is a prerequisite for using habitat models as predictive tools over large areas. The use of a simple sett distribution algorithm circumvents the common problem of subjectively fixing a threshold to discriminate between suitable and unsuitable habitat. In conjunction the models presented here constitute an important contribution to the management of the badger in Denmark and, upon further validation, possibly to similar regions in Northern Europe.     

Confirmation of low genetic diversity and multiple breeding females in a social group of Eurasian badgers from microsatellite and field data

The Eurasian badger (Meles meles) is a facultatively social carnivore that shows only rudimentary co-operative behaviour and a poorly defined social hierarchy. Behavioural evidence and limited genetic data have suggested that more than one female may breed in a social group. We combine pregnancy detection by ultrasound and microsatellite locus scores from a well-studied badger population from Wytham Woods, Oxfordshire, UK, to demonstrate that multiple females reproduce within a social group. We found that at least three of seven potential mothers reproduced in a group that contained 11 reproductive age females and nine offspring. Twelve primers showed variability across the species range and only five of these were variable in Wytham. The microsatellites showed a reduced repeat number, a significantly higher number of nonperfect repeats, and moderate heterozygosity levels in Wytham. The high frequency of imperfect repeats and demographic phenomena might be responsible for the reduced levels of variability observed in the badger.     

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.