Direction of Association between Bite Wounds and Mycobacterium bovis Infection in Badgers: Implications for Transmission

Background

Badgers are involved in the transmission to cattle of bovine tuberculosis (TB), a serious problem for the UK farming industry. Cross-sectional studies have shown an association between bite wounds and TB infection in badgers which may have implications for M. bovis transmission and control, although the sequence of these two events is unclear. Transmission during aggressive encounters could potentially reduce the effectiveness of policies which increase the average range of a badger and thus its opportunities for interaction with other social groups.

Methods

Data were obtained on badgers captured during a long term study at Woodchester Park, UK (1998–2006). Many badgers had multiple observations. At each observation, the badger was assigned a “state” depending on presence of bite wounds and/or TB infection. Hence each badger had a “transition” from the previous state to the current state. We calculated the numbers of each type of transition and the time spent in each state. Transition rates were calculated for each transition category, dividing the number of such transitions by the total time at risk. We compared the rate of bite wound acquisition in infected badgers with that for uninfected badgers and the rate of positive M.bovis test results in bitten badgers with that in unbitten badgers.

Results

The rate of bite wound acquisition in infected badgers (0.291 per year) was 2.09 (95% CI: 1.41, 3.08) times that in uninfected badgers (0.139 per year). The rate of positive M.bovis test results in bitten badgers (0.097 per year) was 2.45 (95% CI: 1.29, 4.65) times that in unbitten badgers (0.040 per year).

Conclusions

We found strong evidence of both potential sequences of events consistent with transmission via bite wounds and distinctive behaviour in infected badgers. The complex relationship between behaviour and infection must be considered when planning TB control strategies.     

Street fighters: Bite force,injury rates,and density of urban Australian water dragons (Intellagama lesueurii)

In an increasingly urbanized world it is imperative that we understand how wildlife responds to this novel anthropogenic landscape, both at the individual‐ and population‐level. Urbanization generally reduces biodiversity, but can also favour particular species and increase their abundance relative to wild populations. When population density increases, so too does the frequency and cost of social interactions. We studied Australian water dragons (Intellagama lesueurii), a species common in urban, semi‐natural and natural areas, to firstly test the prediction that urban populations occur at higher densities, and then determine the consequences of urbanization for combat rates (quantified using wounding) and bite force. We established that urban populations are denser than ones from semi‐natural and natural habitats. We also recorded significantly more wounds in females from urban populations than females from both natural and semi‐natural populations. Urban males also had significantly higher incidence of wounding than males from natural populations. We did not find a difference in male or female bite force among any populations across the urban‐natural gradient. Overall, we found evidence that urbanization results in a higher population density of water dragons and more frequent conspecific combat, but this was not associated with an increase in bite force. These finding suggests that there may be a physiological cost to living in urban habitats related to increased contest rates and wounding.     

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.     

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.     

Body weight as an indication of density-dependent population regulation in badgers (Meles meles) at Woodchester Park, Gloucestershire

The body weights of badgers were examined to look for density-dependent effects of the increase in group size at Woodchester Park, Gloucestershire. The weight of badgers in 21 groups were studied from 1978–1993. A significant negative relationship was found between weight and group size in breeding adult females in autumn and winter, and adult males in summer, such that increase in group size caused a decrease in weight. However, these relationships were affected by year-to-year changes and random variation, and overall there was no downward trend in body weight. The existence of density-dependent effects on badger populations at high density was consistent with previous findings     

To breed or not to breed: an analysis of the social and density-dependent constraints on the fecundity of female badgers (Meles meles).

Data from post-mortem examinations, population density estimates and long term capture-mark-recapture studies have been combined to look at the pattern of reproductive behaviour and the social factors leading to reproductive failure in badgers in Britain. The results are used to evaluate whether the hypothesis that the defence of oestrous females (as opposed to defence of food resources) best explains territorial behaviour and the social organization of badgers. Badgers in Britain have two peaks of reproductive activity, one immediately post partum and one in the summer/autumn. These coincide with two peaks of ovulation, and in the late winter/spring there is a steep rise in the number of sows carrying blastocysts, to reach an asymptote in June for yearling sows and April in older sows. Measured by their contribution to overall productivity, winter/spring matings were much more important than summer/autumn matings, contributing 65% of total autumn blastocysts in yearling sows and 71% of autumn blastocysts in older sows. The relative importance of the two mating periods is reflected in the seasonal pattern of bite wounding in adult male badgers; minor bite wounding in January-March was 2.3 times as frequent as in August-October, and moderate-extensive bite wounding was 3.1 times more frequent. In the populations studied, pre- and post-natal losses were high, with reproductive failure occurring at all stages of the breeding cycle, so that less than 30% of potential productivity was achieved. Indeed 22% of sows failed to develop blastocysts; these had a lower body mass, less body fat, larger adrenal glands, poorer health and higher bite wound scores than sows with blastocysts. Only 44% of adult sows implanted their blastocysts and proceeded to the end of pregnancy. However, it was less easy to identify features characteristic of sows that did or did not go on to implant their blastocysts. Finally, 35% of sows that produced cubs ceased lactation early, and this loss of entire litters was thought to be due to infanticide by dominant sows. The presence of annexe setts correlates with increased productivity in younger sows, and this is thought to be because annexe setts enable younger sows and their cubs to avoid the aggression of older, more dominant sows. Living in large social groups has no net reproductive gain for adult males or females, and there was a decline in productivity (per adult) with increasing group size.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

Dermal Bite Wounds as Indicators of Reproductive Seasonality and Behaviour in the Atlantic Stingray, Dasyatis sabina

Elasmobranch fishes exhibit a series of complex courtship and mating behaviours in which males inflict significant bite wounds to the body of female mates. However, the variety and frequency of mating wounds are not known across a full reproductive season for any species. We examined the distribution and abundance of dermal wounds in adult Atlantic stingrays, Dasyatis sabina, which have a protracted and defined mating season to determine (1) if dermal wounds can be used as indicators of mating activity, (2) whether different categories of bite wounds can be associated with specific mating behaviours, and (3) whether the skin thickness in females is sexually dimorphic. Adults of both sexes showed fresh wounds during the full duration of the mating season (October–June) and there was no relationship between ray size and wound density. Females had more total wounds than males in every month with a maximum average of 20.2 wounds per female in April. Mating wounds were categorized into five distinct forms: single track, double track, bite, margin abrasion and excision. Wounds were randomly distributed over the body of males but concentrated on the posterior half of the disc in females. Each wound type occurred in approximately equal proportions among sexes with the exception of the precopulatory and copulatory-induced margin abrasions which accounted for 13.7% of the total wounds in females but only 3.1% in males. We suggest that the pronounced and concurrent appearance of single track, double track and bite wounds among males results from random premating courtship attacks by males because females cannot be visually discriminated. However, the concentration of wounds on the posterior disc of females is consistent with the possible presence of olfactory cues (e.g. pheromones) that are released at the cloaca. The pectoral fin dermis of females was 50% thicker than that of males, which eliminated the excision of margins during male grasping and functions to reduce female injury. The temporal occurrence of wounds from October through June and peak in April is consistent with previous reproductive studies that show fresh sperm in the reproductive tract of females over the protracted mating period and also ovulation in late March or early April. The importance of social reproductive biting is discussed in relation to the reproductive induction hypothesis proposed to possibly explain the protracted mating of this species. Monitoring of dermal wounds provides a useful non-invasive technique to determine reproductive activity and a means for inference of social relationships in elasmobranch populations.     

Demographic factors associated with prevalence of antibody to Sin Nombre virus in deer mice in the western United States

We used long-term data collected for up to 10 yr (1994-2004) at 23 trapping arrays (i.e., webs and grids) in Arizona, Colorado, Montana, and New Mexico to examine demographic factors known or suspected to be associated with risk of infection with Sin Nombre virus (SNV) in its natural host, the deer mouse (Peromyscus maniculatus). Gender, age (mass), wounds or scars, season, and local relative population densities were statistically associated with the period prevalence of antibody (used as a marker of infection) to SNV in host populations. Nevertheless, antibody prevalence and some of the risk factors associated with antibody prevalence, such as relative population density, gender bias, and prevalence of wounding, varied significantly among sites and even between nearby trapping arrays at a single site. This suggests that local microsite-specific differences play an important role in determining relative risk of infection by SNV in rodents and, consequently, in humans. Deer mouse relative population density varied among sites and was positively and statistically associated with infection prevalence, an association that researchers conducting shorter-term studies failed to demonstrate. Both wounding and antibody prevalence increased with mass class in both males and females; this increase was much more pronounced in males than in females and wounding was more frequent in adult males than in adult females. Prevalence of wounding was greatest among seropositive deer mice, regardless of mass class, but many deer mice without detectable wounds or scars eventually became infected. Many of these patterns, which will be useful in the development of predictive models of disease risk to humans, were only detected through the application of data collected over a long (10-yr) period and with abundant replication.     

Scent-marking with the subcaudal gland by the European badger, Meles meles L.

European badgers, Meles meles, possess a large subcaudal gland which is used for scent-marking the territory and other members of the clan. It was shown experimentally that a badger can distinguish secretions from different individuals. Dominant males scent-mark most, and lactating females more than other females. Scent-marks are distributed along the border of the territory, on bedding materials and around the sett, and on all members of the clan, especially females and cubs. The data support, in many details, the hypothesis that scent-making in badgers serves to establish an asymmetry of contest during encounters with intruders.     

Food habits and habitat selection of suburban badgers (Meles meles) in Japan

A study of the Japanese badger Meles meles anakuma was undertaken in Hinode, a suburb of Tokyo, between 1992 and 1998. Faecal analysis, based on 82 samples, revealed that during spring and summer, earthworms ( Megaseolocidae spp.) occurred at high frequency in the diet, with berries ( Rubus spp.), beetles and persimmon Dymopyrus kaki also eaten during summer months. Scavenged food was eaten in early spring when earthworm availability was low, and badgers switched from worms when persimmon became abundant in autumn. Twenty-one Japanese badgers (14 males and seven females) were radio-tracked. Adult badger home ranges were stable, and those of males [40±19 ( sd ) ha, n =7] were larger than those of females [11±6 ( sd ) ha, n =4]. Badger resting sites in each home range were located within 630 m of each other and categorized as setts or couches. Setts were sited within core areas (30% adaptive kernel method) of home ranges. Most setts were on a sub-ridge and avoided west-facing slopes. Couches, mainly in deciduous forest and forest edge, were generally sited towards the periphery of home ranges. Most badger foods were distributed along ecotones between forestry plantations and farmland; earthworms, their main food from late spring to summer, and berry thickets were both concentrated at the edge of conifer plantations. Persimmon trees, the main food source for badgers in autumn, were also found in agricultural land bordering forest edge. Badger home range size was related to forest edge density.     

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.     

In situ adaptive response to climate and habitat quality variation: spatial and temporal variation in European badger (Meles meles) body weight

Variation in climatic and habitat conditions can affect populations through a variety of mechanisms, and these relationships can act at different temporal and spatial scales. Using post‐mortem badger body weight records from 15 878 individuals captured across the Republic of Ireland (7224 setts across ca. 15 000 km²; 2009–2012), we employed a hierarchical multilevel mixed model to evaluate the effects of climate (rainfall and temperature) and habitat quality (landscape suitability), while controlling for local abundance (unique badgers caught/sett/year). Body weight was affected strongly by temperature across a number of temporal scales (preceding month or season), with badgers being heavier if preceding temperatures (particularly during winter/spring) were warmer than the long‐term seasonal mean. There was less support for rainfall across different temporal scales, although badgers did exhibit heavier weights when greater rainfall occurred one or 2 months prior to capture. Badgers were also heavier in areas with higher landscape habitat quality, modulated by the number of individuals captured per sett, consistent with density‐dependent effects reducing weights. Overall, the mean badger body weight of culled individuals rose during the study period (2009–2012), more so for males than for females. With predicted increases in temperature, and rainfall, augmented by ongoing agricultural land conversion in this region, we project heavier individual badger body weights in the future. Increased body weight has been associated with higher fecundity, recruitment and survival rates in badgers, due to improved food availability and energetic budgets. We thus predict that climate change could increase the badger population across the Republic of Ireland. Nevertheless, we emphasize that, locally, populations could still be vulnerable to extreme weather variability coupled with detrimental agricultural practice, including population management.     

Patterns of wounding in stumptail macaques (macaca arctoides)

A three-year study of the patterns of wounding in a group of stumptail macaques (Macaca arctoides) was conducted at the Yerkes Regional Primate Research Center Field Station. Wounds were classified as punctures, lacerations or abrasions. Data were analyzed to determine if patterns of wounding vary by age/sex class, body part or wound type. Results indicate that adult males receive significantly more total wounds than expected, based on their total time spent in the group. Adult males also receive more serious wounds than other age/sex classes. Low-ranking animals are wounded more often than high-ranking individuals. Moreover, the location of wounds within each age/sex class is non-random. Adult males receive a disproportionate number of wounds on the forequarters, but adult and immature females are wounded disproportionately on the hindquarters. Finally, age/sex classes differ in the number of wounds on individual body parts. Adult males receive more wounds on the head, arms and hands than other age/sex classes, but adult and immature females receive more wounds on the feet than other age/sex classes. These results demonstrate that wounding patterns are clearly non-random and depend on a variety of factors such as age, sex and dominance rank.     

Mating competition and intergroup transfer of males in Tibetan macaques (Macaca thibetana) at Mt. Emei,China

In five groups of seasonally provisioned Tibetan macaques (Macaca thibetana) at Mt. Emei, males were sampled for wounds as an indicator of their competition for females during about 80 days in the 1987 mating season. Quantitative data on intergroup transfer were collected in a period between June 1986 and December 1987. The young adult (YA) males, the most active age-class in mating activity and intergroup transfer, received most of the wounds. Wounds tended to appear more in the front of body for YA and subadults (SA) than they did for middle-old aged (MO) males. This implies that some of the MO males were more active and aggressive in the fights. During the 1.5 year period, 5/6 of the YA and 5/17 of the MO males made intergroup shifts. Although YA males faced a high risk of receiving wounds at transfer, they usually rose in rank. On the other hand, the MO males transferred more smoothly but dropped in rank. The peripheral SA males, which rarely emigrated in the population, were an active component in determining the wounding rate, and the rate and direction of male migration. Three SA immigrants died of severe attacks made by resident males in 1988 and 1991. Adult sex ratios and their variations were considerably reduced with male nonrandom shifts and better conservation of the population.     

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.     

Group size correlates with territory size in European badgers: implications for the resource dispersion hypothesis?

The resource dispersion hypothesis (RDH) predicts that resource heterogeneity can act as a passive cause of group‐living in social carnivores and potentially many other species. One central prediction of the RDH is that territory size and group size are not related, as they are determined by resource dispersion and quality, respectively. In this study we investigated the relationship between territory size, group size and group composition in the European badger, a non‐cooperative social mustelid whose behavioural ecology was central to the development of the RDH. Using data from a long‐term study in the UK, we found that territory size and group size were positively related, contradicting a core prediction of the RDH. Furthermore, territory size was more strongly correlated with the number of adult males in the group than to the number of females or total group size. This result suggests that male badgers may have a more important role in territoriality and receive greater benefits from territory enlargement. These findings are consistent with the predictions of the anti‐kleptogamy hypothesis, and suggest that badger territorial and social behaviour is not purely driven by resource dispersion, but may also be associated with breeding behaviour, as in other mustelids.     

Costs of breeding status in the European badger, Meles meles

Compared with other mammals, reproduction is expected to be particularly costly for European badgers ( Meles meles L.) since both the gestation and mating periods occur in the winter when the animals are inactive and feed little. This paper assesses the costs of breeding status in both male and female badgers, in terms of body condition, ectoparasite load, haematology and mortality. By the end of the lactation period (May), breeding females had suffered a marked loss in body condition, although there was no such cost associated with gestation. However, females regained the weight that they had lost by the autumn following lactation. Lactating two-year-olds experienced a higher age-specific mortality than those that did not breed, but there was no difference among older females. No costs of breeding status could be detected among males at the end of the spring mating period. However, breeding males sustained testicular activity later into the summer than non-breeders, and by the autumn they had acquired more bite wounds and become anaemic. This suggests that there was a physiological cost associated with extended testicular activity in breeding males.     

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.     

苏格兰拉姆岛上野化山羊种群的取食生态学

2000年6～11月对苏格兰拉姆岛上野化山羊(Capra hircus)种群的取食生态学进行了研究.研究表明:山羊的觅食回合长度变化范围从1min到460 min不等,平均觅食回合长度是103.1±15.0 (SD)min,雌性动物的觅食回合长度较雄性的长(P=0.077).野化山羊单位时间的取食频率平均为46.3±0.6 口/min,取食频率随性别(P=0.023)和月份(P<0.001)而显著变化.雄性山羊在繁殖交配之前(6～7月)和之后(10～11月)的取食频率比繁殖交配期中(9～10月)的快(P<0.008),但雌性动物并没有这样的变化(P=0.327).雄性动物在繁殖交配期中的取食时间显著减少.雌、雄两性动物在取食频次和取食时间方面的这些差异可能导致该山羊种群在食物摄入量上的性别差异:雌性山羊的食物摄入量相对比较稳定,而雄性山羊的摄入量则变动很大.估计的食物摄入量随月份而下降(尽管9月份以后有一微小幅度的上升),这意味着拉姆岛上的山羊种群在食物匮乏而天气寒冷潮湿的冬季可能面临着能量收支不平衡的威胁.