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1. Experimental laboratory evidence suggests that animals with disrupted social systems express weakened relationship strengths and have more exclusive social associations, and that these changes have functional consequences. A key question is whether anthropogenic pressures have a similar impact on the social structure of wild animal communities. 2. We addressed this question by constructing a social network from 6 years of systematically collected photographic capture–recapture data spanning 1,139 individual adult female Masai giraffes inhabiting a large, unfenced, heterogeneous landscape in northern Tanzania. We then used the social network to identify distinct social communities, and tested whether social or anthropogenic and other environmental factors predicted differences in social structure among these communities. 3. We reveal that giraffes have a multilevel social structure. Local preferences in associations among individuals scale up to a number of distinct, but spatially overlapping, social communities, that can be viewed as a large interconnected metapopulation. We then find that communities that are closer to traditional compounds of Indigenous Masai people express weaker relationship strengths and the giraffes in these communities are more exclusive in their associations. 4. The patterns we characterize in response to proximity to humans reflect the predictions of disrupted social systems. Near bomas, fuelwood cutting can reduce food resources, and groups of giraffes are more likely to encounter livestock and humans on foot, thus disrupting the social associations among group members. Our results suggest that human presence could potentially be playing an important role in determining the conservation future of this megaherbivore. published

Studies increasingly show that social connectedness plays a key role in determining survival, in addition to natural and anthropogenic environmental factors. Few studies, however, integrated social, non-social and demographic data to elucidate what components of an animal's socio-ecological environment are most important to their survival. Female giraffes ( Giraffa camelopardalis ) form structured societies with highly dynamic group membership but stable long-term associations. We examined the relative contributions of sociability (relationship strength, gregariousness and betweenness), together with those of the natural (food sources and vegetation types) and anthropogenic environment (distance from human settlements), to adult female giraffe survival. We tested predictions about the influence of sociability and natural and human factors at two social levels: the individual and the social community. Survival was primarily driven by individual- rather than community-level social factors. Gregariousness (the number of other females each individual was observed with on average) was most important in explaining variation in female adult survival, more than other social traits and any natural or anthropogenic environmental factors. For adult female giraffes, grouping with more other females, even as group membership frequently changes, is correlated with better survival, and this sociability appears to be more important than several attributes of their non-social environment.

Background Appropriate social interactions influence animal fitness by impacting several processes, such as mating, territory defense, and offspring care. Many studies shedding light on the neurobiological underpinnings of social behavior have focused on nonapeptides (vasopressin, oxytocin, and homologues) and on sexual or parent-offspring interactions. Furthermore, animals have been studied under artificial laboratory conditions, where the consequences of behavioral responses may not be as critical as when expressed under natural environments, therefore obscuring certain physiological responses. We used automated recording of social interactions of wild house mice outside of the breeding season to detect individuals at both tails of a distribution of egocentric network sizes (characterized by number of different partners encountered per day). We then used RNA-seq to perform an unbiased assessment of neural differences in gene expression in the prefrontal cortex, the hippocampus and the hypothalamus between these mice with naturally occurring extreme differences in social network size. Results We found that the neurogenomic pathways associated with having extreme social network sizes differed between the sexes. In females, hundreds of genes were differentially expressed between animals with small and large social network sizes, whereas in males very few were. In males, X-chromosome inactivation pathways in the prefrontal cortex were the ones that better differentiated animals with small from those with large social network sizes animals. In females, animals with small network size showed up-regulation of dopaminergic production and transport pathways in the hypothalamus. Additionally, in females, extracellular matrix deposition on hippocampal neurons was higher in individuals with small relative to large social network size. Conclusions Studying neural substrates of natural variation in social behavior in traditional model organisms in their habitat can open new targets of research for understanding variation in social behavior in other taxa.

Natural disasters can cause rapid demographic changes that disturb the social structure of a population as individuals may lose connections. These changes also have indirect effects as survivors alter their within-group connections or move between groups. As group membership and network position may influence individual fitness, indirect effects may affect how individuals and populations recover from catastrophic events. Here we study changes in the social structure after a large predation event in a population of wild house mice (Mus musculus domesticus), when a third of adults were lost. Using social network analysis, we examine how heterogeneity in sociality results in varied responses to losing connections. We then investigate how these differences influence the overall network structure. An individual's reaction to losing associates depended on its sociality prior to the event. Those that were less social before formed more weak connections afterwards, while more social individuals reduced the number of survivors they associated with. Otherwise, the number and size of social groups were highly robust. This indicates that social preferences can drive how individuals adjust their social behaviour after catastrophic turnover events, despite the population's resilience in social structure., Proceedings of the Royal Society B: Biological Sciences, 287 (1926), ISSN:0080-4649, ISSN:0950-1193, ISSN:1471-2954, ISSN:0962-8452

Social interactions are critically important for survival and impact overall-health, but also impose costs on animals, such as exposure to contagious agents. The immune system can play a critical role in modulating social behavior when animals are sick, as has been demonstrated within the context of “sickness behaviors.” Can immune molecules affect or be affected by social interactions even when animals are not sick, therefore serving a role in mediating pathogen exposure? We tested whether markers of immune function in both the blood and the brain are associated with gregariousness, quantified as number of animals interacted with per day. To do this, we used remote tracking of social interactions of a wild population of house mice (Mus musculus domesticus) to categorize animals in terms of gregariousness. Blood, hair, brain and other tissue samples from animals with extreme gregariousness phenotypes were collected. We then tested whether the levels of three important cytokines (TNF-α, IFN-γ and IL-1β) in the serum, cortex and hypothalamus of these animals could be explained by the gregariousness phenotype and/or sex of the mice. Using the hair as a long-term quantification of steroid hormones, we also tested whether corticosterone, progesterone and testosterone differed by social phenotype. We found main effects of gregariousness and sex on the serum levels of TNF-α, but not on IFN-γ or IL-1β. Brain gene expression levels were not different between phenotypes. All hair steroids tended to be elevated in animals of high gregariousness phenotype, independent of sex. In sum, elements of the immune system may be associated with gregariousness, even outside of major disease events. These results extend our knowledge of the role that immune signals have in contributing to the regulation of social behaviors outside periods of illness.

Associating with relatives in social groups can bring benefits such as reduced risk of aggression and increased likelihood of cooperation. Competition among relatives over limited resources, on the other hand, can induce individuals to alter their patterns of association. Population density might further affect the costs and benefits of associating with relatives by altering resource competition or by changing the structure of social groups; preventing easy association with relatives. Consequently, the overlap between genetic and social structure is expected to decrease with increasing population size, as well as during times of increased breeding activity. Here, we use multi-layer network techniques to quantify the similarity between long-term, high resolution genetic, and behavioral data from a large population of free-ranging house mice (Mus musculus domesticus), studied over 10 years. We infer how the benefit of associating with genetically similar individuals might fluctuate in relation to breeding behavior and environmental conditions. We found a clear seasonal effect, with decreased overlap between social and genetic structure during summer months, characterized by high temperatures and high breeding activity. Though the effect of overall population size was relatively weak, we found a clear decrease in the overlap between genetic similarity and social associations within larger groups. As well as longer-term within-group changes, these results reveal population-wide short-term shifts in how individuals associate with relatives. Our study suggests that resource competition modifies the trade-off between the costs and benefits of interacting with relatives.

Endocrine data from wild populations provide important insight into social systems. However, obtaining samples for traditional methods involves capture and restraint of animals, and/or pain, which can influence the animal’s stress level, and thereby undesirable release of hormones. Here, we measured corticosterone, testosterone and progesterone in the hair of 482 wild-derived house mice that experienced sexual competition while living under semi-natural conditions. We tested whether sex, age, weight and indicators of sexual maturity, reproduction and social conflicts predict hormone concentrations measured in hair (sampling at endpoint). We show that body weight, sex and age significantly predict cumulative testosterone and progesterone levels, allowing the differentiation between subadults and adults in both sexes. Corticosterone was only slightly elevated in older males compared to older females and increased with the level of visible injuries or scars. Testosterone in males positively correlated with body weight, age, testes size, and sperm number. Progesterone in females significantly increased with age, body weight, and the number of embryos implanted throughout life, but not with the number of litters when controlled for age and weight. Our results highlight the biological validity of hair steroid measurements and provide important insight into reproductive competition in wild house mice.

Background Communal nursing in house mice is an example of cooperation where females pool litters in the same nest and indiscriminately nurse own and other offspring despite potential exploitation. The direct fitness benefits associated with communal nursing shown in laboratory studies suggest it to be a selected component of female house mice reproductive behaviour. However, past studies on communal nursing in free-living populations have debated whether it is a consequence of sharing the same nest or an active choice. Here using data from a long-term study of free-living, wild house mice we investigated individual nursing decisions and determined what factors influenced a female’s decision to nurse communally. Results Females chose to nurse solitarily more often than expected by chance, but the likelihood of nursing solitarily decreased when females had more partners available. While finding no influence of pairwise relatedness on partner choice, we observed that females shared their social environment with genetically similar individuals, suggesting a female’s home area consisted of related females, possibly facilitating the evolution of cooperation. Within such a home area females were more likely to nest communally when the general relatedness of her available options was relatively high. Females formed communal nests with females that were familiar through previous associations and had young pups of usually less than 5 days old. Conclusions Our findings suggest that communal nursing was not a by-product of sharing the same nesting sites, but females choose communal nursing partners from a group of genetically similar females, and ultimately the decision may then depend on the pool of options available. Social partner choice proved to be an integrated part of cooperation among females, and might allow females to reduce the conflict over number of offspring in a communal nest and milk investment towards own and other offspring. We suggest that social partner choice may be a general mechanism to stabilize costly cooperation. Electronic supplementary material The online version of this article (10.1186/s12983-018-0251-3) contains supplementary material, which is available to authorized users.

Abiotic, biotic and human influences are factors that can affect animal home ranges. We calculated home range sizes of adult giraffes in the Tarangire-Manyara region of northern Tanzania (N = 132 giraffes with data collected over 6 years), and investigated correlations between home range sizes and environmental and anthropogenic factors (for a subset of N = 71 giraffes). We used a 95% kernel utilization distribution to define home ranges and modelled home range size as a function of environmental and anthropogenic predictors using multiple linear regression and model selection. We also computed home range sizes of giraffes using 100% minimum convex polygons to compare with estimates from previously published studies, and tested the relationship between rainfall and home range sizes of giraffes across Africa. Average kernel home range sizes were 114.6 km2 for females (N = 109) and 157.2 km2 for males (N = 23). Adult female giraffe home range sizes (N = 67) were negatively correlated with distance to densely populated towns. Females living closer to towns had significantly larger home ranges, suggesting a need to range farther to avoid human-impacted areas while obtaining critical resources. No such relationship was evident with bomas, which are homesteads built by indigenous pastoralist people, suggesting that female giraffes are tolerant of traditional land uses. Mean annual rainfall explained 74% of the variation in home range sizes of giraffes across the African continent, with smaller home ranges in regions with higher rainfall and thus greater productivity, providing additional evidence that access to critical resources mediates home range size of this megaherbivore. Quantifying home range sizes and identifying ecological and anthropogenic factors affecting space use can provide insights into mechanisms driving use of space and help wildlife managers make informed decisions that improve conservation plans for at-risk species such as giraffes.

Fission–fusion dynamics hypothetically enable animals to exploit dispersed and ephemeral food resources while minimizing predation risk. Disentangling factors affecting group size and composition of fission–fusion species facilitates their management and conservation. We used a 6-year data set of 2888 group formations of Masai giraffes in Tanzania to investigate determinants of social group size and structure. We tested whether ecological (lion density, vegetation structure, and prevalence of primary forage plants), anthropogenic (proximity to human settlements), temporal (rainy or dry season), and social (local giraffe density, adult sex ratio, and proportion of calves) factors explained variation in group size and sex- and age-class composition. Food availability rather than predation risk mediated grouping dynamics of adult giraffes, while predation risk was the most important factor influencing congregations with calves. Smallest group sizes occurred during the food-limiting dry season. Where predation risk was greatest, groups with calves were in bushlands more than in open grasslands, but the groups were smaller in size, suggesting mothers adopted a strategy of hiding calves rather than a predator-detection-and-dilution strategy. Groups with calves also were farther from towns but closer to traditional human compounds (bomas). This may be due to lower predator densities, and thus reduced calf predation risk, near bomas but higher human disturbance near towns. Sex- and age-based differences in habitat use reflected nursing mothers’ need for high-quality forage while also protecting their young from predation. Our results have implications for conservation and management of giraffes and other large-bodied, herd-forming ungulates in heterogeneous environments subject to anthropogenic threats.

International audience; Individual recognition, the ability to discriminate between members of a social group according to their distinctive characteristics, is a sophisticated form of social recognition. Several laboratory studies demonstrated individual recognition in rodents, but not under natural conditions. We combined behavioral observations and an experiment to assess individual recognition in free-living African striped mice Rhabdomys pumilio. Striped mice live in groups that share a nest, but forage alone during the day. Interactions among group members are typically amicable and take place at the nest. In order to determine whether striped mice have individualized relationships, we analyzed whether focal mice interacted repeatedly more often with one specific group member. Data from 58 focal individuals from 15 groups revealed that the individual with which amicable interactions occurred the most was often alone in one mode of the data distribution, indicating that behavior toward this specific individual differed compared to the behavior to all other group members. Striped mice consistently preferred this specific individual during the following weeks. Preferred social partners of both breeders and younger non-breeders were young non-breeders. Sex-specific effects were obtained only for young adult males and breeding females, which both associated more often with the opposite sex. For 11 young adults from 11 groups, the preferred social partner was trapped and removed, either overnight (experiment) or only during the day (control), when striped mice forage solitarily and usually do not interact with group members. Striped mice interacted on average more with their preferred social partner after it returned in the experimental group compared to the control (7 of 11 experiments), but the difference was not significant (P = 0.16). Our study indicates that striped mice might have individualized relationships, although they failed to significantly indicate individual recognition during experimental removals. Future studies will have to test the adaptive function of individualized relationships in African striped mice

When infected, animals change their behaviors in several ways, including by decreasing their activity, their food and water intake, and their interest in social interactions. These behavioral alterations are collectively called sickness behaviors and, for several decades, the main hypotheses put forward to explain this phenomenon were that engaging in sickness behaviors facilitated the fever response and improved the likelihood of host survival. However, a new hypothesis was recently proposed suggesting that engaging in sickness behaviors may serve to protect kin. We tested this kin protection hypothesis by combining a field and a laboratory experiment in house mice. In both experiments, we induced sickness behaviors by administration of a pro-inflammatory agent. In the field experiment, we then collected genetic data and assessed whether relatedness affected the intensity of sickness behaviors. In the lab experiment, we manipulated relatedness in small social groups and assessed whether having a closely related individual (a sibling) in the group altered social interactions or visits to common resources (such as food and water containers) once immune-challenged. Our results do not support the kinship protection hypothesis and therefore advance our understanding of why such an apparently costly set of behavioral changes would be evolutionarily maintained., Scientific Reports, 8 (1), ISSN:2045-2322

In group living animals, reproductive competition plays an important role in shaping social relationships and associations among female group members. In this study, we investigated the impact of male presence on the development of female-female competition and female sociality in groups of female wild house mice, using physiological and behavioral parameters. We predicted that, by eliciting intra-sexual competition, males influence social relationships among female group members and thus affect female associations to potential cooperation partners. To test this hypothesis we compared stress hormone production, the frequency of agonistic interactions, social hierarchies and social partner preferences in groups of unrelated, unfamiliar females in the absence and presence of males. Our results revealed no indication that the introduction of males into all-female groups of wild house mice elicited increased competition among female group members, neither on the physiological nor on the behavioral level. We found no effect of male presence on female glucocorticoid secretion, aggression, dominance hierarchies or on the females' sociability. Females thus seem not to intensely compete over access to males. This female ability to behaviorally and physiologically deal with even previously unfamiliar same-sex group members may be an important feature of female house mouse societies. In fact, it could be a necessary prerequisite to establish cooperative relationships between females in the context of reproduction, such as communal nursing of young.

International audience; An individual's home range determines its access to resources, significantly influencing its fitness. Food availability and population density are considered to be among the primary factors influencing home range sizes; however, no study has experimentally tested whether these two factors affect home range sizes independently. This is important as these two factors correlate significantly with each other, making it difficult to differentiate the effect of one from the other. First, we supplemented food to 23 female African striped mice, Rhabdomys pumilio, belonging to 15 different groups. To avoid an increase in population density by immigration, we also provided food to neighbouring groups. Although population density did not increase, female home range sizes decreased by 43.1%. In a second experiment, we manipulated population density by removing entire social groups of striped mice. We carried out experiments within 7 weeks, a period short enough to control for a change in natural food availability. Experimental decrease of population density caused an increase of female home range sizes of 44.3%. The degree of home range overlap between female striped mice was unaffected by supplemental feeding or by reduction of population density. However, female home range sizes were negatively affected by the total number of female neighbours, especially of heavier individuals. In addition, after removal, females significantly decreased their overlap with neighbouring breeding females indicating that competition with same-sex individuals shapes females' home ranges. This is the first comprehensive experimental field study demonstrating that an increase in food availability and a decrease in population density independently affect home range sizes of individuals.

The t haplotype in house mice is a well-known selfish genetic element with detrimental, nonadditive fitness consequences to its carriers: recessive lethal mutations cause t/t homozygotes to perish in utero. Given the severe genetic incompatibility imposed by the t haplotype, we predict females to avoid fertilization by t haplotype incompatible males. Indeed, some of the strongest evidence for compatibility mate choice is related to the t haplotype in house mice. However, all previous evidence for compatibility mate choice in this system is based on olfactory preference. It is so far unknown how general these preferences are and whether they are relevant in an actual mating context. Here, we assess female compatibility mate choice related to t haplotypes in a setting that – for the first time – allowed females to directly interact and mate with males. This approach enabled us to analyse female behaviour during the testing period, and the resulting paternity success and fitness consequences of a given choice. We show that genetic incompatibilities arising from the t haplotype had severe indirect fitness consequences and t females avoided fertilization by t incompatible males. The results are inconclusive whether this avoidance of t fertilization by t females was caused by pre- or post-copulatory processes.

We studied the feeding ecology of the bongo (Tragelaphus eurycerus) in the Dzanga National Park in the rainforest of the Central African Republic to understand better the trade-off between food selectivity, ranging behaviour and social organization of a large, forest-dwelling, social antelope. Food plants and vegetation types were registered along a 311-km route travelled by bongos. Food availability was determined by identifying and counting the plants in 19 randomly chosen forest plots. Bongos showed pronounced selectivity for 26 out of 100 woody forest species. They predominantly consumed younger leaves, which suggests that high protein and low fibre content influence plant choice. In addition to leaves, bongos also ate fruits of two and flowers of one species. Furthermore, the diet was supplemented by grasses and herbs consumed on large natural licks. Such licks were regularly visited by the bongos. According to Jarman's ecological classification of antelopes, selective browsers are relatively small and live alone or in pairs to avoid competition over food. The bongo's large size and gregariousness should not allow it to survive in the rainforest as a pure selective browser. Our data suggest that the bongo relies on the opportunity to graze in bulk which it finds on the natural licks. We hypothesize that such licks either limit the distribution of bongos in other rainforest areas or allow larger group sizes than in areas without licks.

Oxytocin manipulation has been implicated in the facilitation of social and cooperative behaviours, either through increasing positive and cooperative social interactions, or facilitating bond formation. Here we aimed to determine whether peripheral administration of oxytocin would affect the propensity of unrelated female house mice, Mus musculus domesticus, to cooperate. To investigate this, we used female house mice, with their cooperative ability to communally nurse offspring. Pairs of unfamiliar females received intraperitoneal injections of oxytocin over a 3-day cohabitation period. Following this initial phase, a male was introduced and allowed to mate with the females. We monitored how long it took females to establish and successfully cooperate in the raising of a communal litter. Oxytocin did not affect the females' ability to reproduce. However, oxytocin-treated females took significantly longer to establish a successful communal litter (with pups of both partners being weaned) than saline-treated control females. This delay in communal nursing was due to higher pup mortality and loss of first-born litters in the oxytocin group during their first reproductive event. We conclude that administration of exogenous oxytocin during the early stages of the female relationship delayed the tendency of female house mice to affiliate and cooperate in rearing a communal litter. Our findings contribute to the growing field of oxytocin-based studies and sheds light on the potential long-term effects of oxytocin during early pairwise social interactions.

Punishment is claimed as an important mechanism to stabilise costly cooperation in humans, but its importance in social animals has been questioned recently due to both conceptual considerations and a lack of empirical evidence (only few published studies). We empirically tested whether there is evidence for punishment in communally nursing house mice (Mus musculus domesticus, direct descendants of “wild” animals). Communally breeding females pool their litters and raise all offspring together, indiscriminately caring for own and other offspring. Such a situation resembles a public good and provides scope for exploitation if females vary in their relative contributions to the joint nest (offspring number). We allowed two females to communally breed and conducted removal experiments both in the presence and absence of pups. We aimed to test whether reduced investment by one of the females (induced through separation from the partner and their combined offspring for 4 or 12 hours) leads to increased aggression by the other female after the reunion. We found no evidence for punishment, on the contrary, females increased socio-positive behaviours. The costs of losing a partner in a communally breeding species might be too high and hinder the evolution of punishment. Our findings add to a growing list of examples questioning the role of punishment in cooperating non-human animals and emphasise the importance of empirical testing of its assumptions and predictions.

Among species providing uniparental care, the caring parent faces time constraints and may have to compromise offspring care/protection for self-maintenance. In most mammalian species females raise their offspring without receiving help from males. Communal nesting, when multiple females share a single nest where they rear their pups together, may have evolved as a mutually beneficial cooperative behaviour to reduce mothers' nest attendance without increasing the time their offspring are left alone. We tested this hypothesis using data collected in a free-living house mouse, Mus musculus domesticus, population in which reproduction occurred in nestboxes and was closely monitored. Individuals were fitted with transponders allowing automatic recording of their location, and a genetic parentage analysis confirmed maternal identity. Compared with mothers raising their pups solitarily, communally nesting mothers spent less time inside their nest. Their pups, however, were left alone for a similar amount of time as solitarily raised pups. The time communal litters were left alone did not covary with the kinship of communally nesting females. These results indicate that communally nesting mothers can allocate more time to foraging or territorial defence without impairing the amount of maternal attention received by their offspring. Nevertheless, communally nesting mothers showed some overlap in their stays at the nest. Offspring may benefit from more regular meals while mothers may gain information on the partner's contribution to combined maternal care which could potentially prevent cheating.

In many species, females have evolved behavioral strategies to reduce the risk of infanticide. For instance, polyandry can create paternity confusion that inhibits males from killing offspring they could have sired. Here, the authors propose that females could socially obtain the same benefits by nesting communally. Singly sired litters could be perceived as a large multiply sired litter once pooled together in a single nest. Long-term data from a wild house mouse population showed that monandrous litters (singly sired) were more common in communal than in solitary nests and 85% of them were raised with litters sired by different males hence becoming effectively polyandrous (multiply sired). These socially polyandrous litters had significantly higher offspring survival than genetically or socially monandrous litters and reached a similar survival to that of multiply sired litters raised in solitary or communal nests. Furthermore, the number of sires within nests significantly improved offspring survival whereas the number of mothers did not. These results suggest that the survival benefits associated with communal nesting are driven by polyandry and not communal defense. This socially mediated polyandry was as efficient as multiple paternity in preventing infanticide, and may also occur in other infanticidal and polytocous species where the caring parent exhibits social behavior., Lay Summary Communal nesting allows social females to avoid infanticide in much the same way as multiple mating: through paternity confusion. Litters sired by a single male can be perceived as a large multiply-sired litter once pooled together in a communal nest. This socially mediated polyandry was as efficient as multiple paternity in preventing infanticide. The number of mothers in communal nests did not explain offspring survival hence rejecting the communal defense hypothesis.

Communal nursing, the provision of milk to non-offspring, has been argued to be a non-adaptive by-product of group living. We used 2 years of field data from a wild house mouse population to investigate this question. Communal nursing never occurred among females that previously lacked overlap in nest box use. Females nursed communally in only 33 % of cases in which there was a communal nursing partner available from the same social group. Solitarily nursing females were not socially isolated in their group; nevertheless, high spatial associations prior to reproduction predict which potential female partner was chosen for communal nursing. An increase in partner availability increased the probability of communal nursing, but population density itself had a negative effect, which may reflect increased female reproductive competition during summer. These results argue that females are selective in their choice of nursing partners and provide further support that communal nursing with the right partner is adaptive. Electronic supplementary material The online version of this article (doi:10.1007/s00114-013-1130-6) contains supplementary material, which is available to authorized users.

The animals primarily infected by Francisella tularensis are rapidly consumed by scavengers, hindering ecologic investigation of the bacterium. We describe a 2012 natural tularemia epizootic among house mice in Switzerland and the assessment of infection of exposed humans. The humans were not infected, but the epizootic coincided with increased reports of human cases in the area.

Infection may modify the behaviour of the host and of its conspecifics in a group, potentially altering social connectivity. Because many infectious diseases are transmitted through social contact, social connectivity changes can impact transmission dynamics. Previous approaches to understanding disease transmission dynamics in wild populations were limited in their ability to disentangle different factors that determine the outcome of disease outbreaks. Here we ask how social connectivity is affected by infection and how this relationship impacts disease transmission dynamics. We experimentally manipulated disease status of wild house mice using an immune challenge and monitored social interactions within this free-living population before and after manipulation using automated tracking. The immune-challenged animals showed reduced connectivity to their social groups, which happened as a function of their own behaviour, rather than through conspecific avoidance. We incorporated these disease-induced changes of social connectivity among individuals into models of disease outbreaks over the empirically-derived networks. The models revealed that changes in host behaviour frequently resulted in the disease being contained to very few animals, as opposed to becoming widespread. Our results highlight the importance of considering the role that behavioural alterations during infection can have on social dynamics when evaluating the potential for disease outbreaks.

Conditional adjustment of cooperativeness to the expected pay-off might be a useful strategy to avoid being exploited in public good situations. Parental care provided towards all offspring in a communal nest (containing offspring of several females) resembles a public good. Females indiscriminately caring for all young share the costs equally, but the pay-off may vary depending on their contribution to the joint nest (number of own offspring). Females with fewer offspring in the joint nest will be exploited and overinvest relative to their contribution. We experimentally created a situation of high conflict in communally nursing house mice, by using a genetic tool to create a difference in birth litter sizes. Females in the high conflict situation (unequal litter sizes at birth) showed a reduced propensity to give birth as part of a communal nest, therefore adjusting their cooperativeness to the circumstances.

Social interactions are critical for reproduction in many animals. Since several pathogens are transmitted by social contact, females searching for mating partners should select males that can signal being healthy. Not all signals, however, may be reliable, since males from a number of species can overcome behavioural symptoms of infection when mating opportunities are available. Here, we manipulated sickness status of male house mice, Mus musculus domesticus, by administering an immune challenge (lipopolysaccharide, LPS) and studied the consequences of this manipulation for two signals that function in mate attraction in this species: ultrasonic vocalizations and darcin (a urinary protein). Additionally, we quantified female visits to immune-challenged and control males, and the males' plasma testosterone levels. LPS-injected males had lower darcin and lower regular ultrasonic syllable production than control-injected males, while producing a larger number of high-frequency ultrasonic syllables. We conclude that immune-challenged male mice presented with a receptive female cannot maintain the production of sexually attractive signals. Females might use some of these cues when making mating decisions, since they spent significantly less time near LPS-injected males. Testosterone was reduced in LPS-injected males and could be a unifying mechanism downregulating both of the traits quantified. Darcin and ultrasonic vocalizations produced in the context of courtship may therefore function as reliable indicators of current health status.

Our research focuses on mechanisms that promote and stabilize social behavior, fitness consequences of cooperation, and how interactions with conspecifics structure groups and populations. To this end, we studied wild house mice (Mus musculus domesticus) in the laboratory, in semi-natural enclosures and in the field. In 2002, we initiated a project on a free-living population of house mice in a barn near Zurich, Switzerland, where mice were equipped with RFID transponders and were provided with 40 nest boxes for resting and breeding. The population typically comprised between 250 and 400 mice. To analyze social group membership, social interactions and social preferences of the mice in our study population over their lifespan, we installed a continuous transponder reading system (AniLoc, FBI Science GmbH, Germany). Mice accessed nest boxes through tunnels equipped with two antennas each. When a mouse implanted with an RFID transponder passed the electromagnetic field of an antenna, its identity was transmitted and registered in real time with AniLoc. Additionally, body weights of mice were automatically registered at eight drinking facilities (Intelliscale, FBI Science GmbH, Germany). Here, a mouse sits on a freely movable platform that connects to a scale registering body weight when drinking, and an antenna around the head of the water bottle registers the drinking individual’s RFID transponder. The system enabled continuous remote monitoring of the behavior of a free-living, open population of house mice, when using nest boxes and when drinking. Since such safe places are an important resource for survival and reproduction, time of day, duration and frequency of meetings with conspecifics reveal information about the function of their interactions. Trigger efficiency of antennas was 98.2 %. Mice entered and left the nest boxes with an average speed of 0.03 m/s, which is within the antennas’ detection capacity (detection speed of 1 m/s or 3.6 km/h). The antenna devices documented not only social structuring of our study population but also spatial genetic structuring. The observation that mice lived in rather closed social groups and tended to share nest boxes with relatives highlights the importance of kin selection for the evolution and maintenance of social behavior. We suggest that such automatic recording of activity, spatial distribution and social interactions is helpful not only in field studies, for a variety of species, but also in captivity or laboratory studies, to answer basic questions in behavioral ecology, population ecology, population genetics, conservation biology, disease ecology, or animal welfare.

Information about the welfare and husbandry of pet and laboratory fish is scarce although millions of fish are sold in pet shops and used in laboratory research every year. Inadequate housing conditions can cause behavioural problems also in fish since they are complex animals with sophisticated behaviour. In this study, we investigated the influence of environmental complexity on compartment preference and behaviour in zebrafish (Danio rerio) and checker barbs (Puntius oligolepis). For the preference test, large aquaria were divided by two semi-transparent walls of Plexiglas into an empty compartment, a structured compartment enriched with plants and clay pots, and a smaller compartment in-between, where food was provided. For observation, the empty and structured compartments were divided into six zones of similar size by defining three vertical layers and two horizontal areas (back vs. front area). Seven groups of six to nine zebrafish and seven groups of seven or eight checker barbs were observed on four days each (within a time period of ten days) to assess compartment use and activity, and to assess behavioural diversity and use of zones within compartments. Both zebrafish and checker barbs showed a significant preference for the structured compartment. Nevertheless, in neither species did behavioural diversity differ between the empty and structured compartment. Zebrafish used all zones in both compartments to the same extent. Checker barbs, however, used the structured compartment more evenly than the empty compartment, where they mainly used the lower and middle zones. These results suggest that zebrafish and checker barbs have a preference for complex environments. Furthermore, they indicate that the behavioural and ecological needs of fish may vary depending on species, and recommendations for husbandry should be specified at species level.

In captive carnivores, species-specific behaviour is often restricted by inadequate feeding regimens. Feeding live prey is not feasible in most places and food delivery is often highly predictable in space and time which is considerably different from the situation in the wild. As a result, captive carnivores are often inactive, show little behavioural diversity and are prone to behavioural problems such as stereotypic pacing. Using artificial feeding devices to substitute for natural food resources is a way to address these problems. In a group of four red fox ( Vulpes vulpes ), we compared a conventional feeding method to four different methods through the use of feeding enrichment that were based on natural foraging strategies of opportunistic carnivores. Feeding enrichments consisted of electronic feeders delivering food unpredictable in time which were successively combined with one of the three additional treatments: a self-service food box (allowing control over access to food), manually scattering food (unpredictable in space), and an electronic dispenser delivering food unpredictably both in space and time. The aim of administering feeding enrichment in this study was to stimulate appetitive (food searching) behaviour and to increase time spent in feeding. Compared to conventional feeding, diversity of behaviour and overall activity were significantly enhanced in the presence of electronic feeders in all four foxes (EF>CON1=CON2, EF+SF>CON1=CON2, EF+MS>CON1=CON2, EF+ED>CON1=CON2). Behavioural diversity was highest when the foxes had control over access to food (EF+SF), while the manual scattering of food (EF+MS) and the electronic dispenser (EF+ED) enhanced food searching behaviour. These results indicate that in opportunistic carnivores natural foraging and feeding behaviour can be stimulated by simple feeding enrichment strategies, and that foraging behaviour is stimulated most when food delivery is unpredictable both in space and time.

The relative plasticity hypothesis predicts that alternative tactics are associated with changes in steroid hormone levels. In species with alternative male reproductive tactics, the highest androgen levels have usually been reported in dominant males. However, in sociable species, dominant males show amicable behaviors to gain access to females, which might conflict with high testosterone levels. We compared testosterone, corticosterone, and resting metabolic rate in male striped mice (Rhabdomys pumilio) following a conditional strategy with three different reproductive tactics: (i) philopatric group-living males, (ii) solitary-living roamers, (iii) dominant but sociable group-living territorial breeders. Philopatrics had the lowest testosterone but highest corticosterone levels, suggesting that they make the best of a bad job. Dominant territorial breeders had lower testosterone levels than roamers, which have a lower competitive status. Roamers had the highest testosterone levels, which might promote risky behavior, such as invading territories defended by territorial males. Roamers also had lower resting metabolic rates than either type of group-living males. Our results suggest that dominant males' testosterone levels reflect a trade-off between low testosterone amicable behavior and high testosterone dominance behavior.

Female house mice (Mus domesticus) are known to perform communal breeding more often with kin than with non-kin partners. When mice are grouped in semi-natural enclosures, related females develop pre- reproductive alliances more frequently than unrelated ones. However, little is known about the behavioural mechanisms and the factors promoting kin preferential cooperative associations in female mice. Here we evaluate the relative importance of familiarity and age disparity on the pre-reproductive development of agonistic behaviour and spatial associations within groups of three related females, freely interacting in semi-natural indoor enclo- sures. We found that familiarity clearly promoted female alliance formation by reducing aggression and enhancing spatial cohabitation, while genetic relatedness per se did not alleviate the effects of unfamiliarity on female grouping. Older sisters clearly dominated younger ones, even if they had lower body weights, or if they had to confront two allied littermate sisters. Hence, our findings corroborate previous findings on familiarity as a proxy for kin-preferential alliance formation in female mice. More- over, we observed a strong impact of age stratification on female aggression-mediated dominance development. We suggest that this age effect could be expected from a queuing-for-reproduction view of younger house mouse females.

The presence of selfish genetic elements can have fatal consequences for populations that harbor them. In the well known t haplotype in wild house mice, large proportions of the population die from t/t recessive lethal effects. Due to strong advantages at the gamete level (drive), t haplotypes nevertheless occur at substantial frequencies. The stable presence of a lethal is not the only effect of the t. It also distorts the fate of mutations that differentially affect male and female survival and reproduction (such as in sexual conflict), by giving male selective effects a strong advantage over female selective effects. In a recent study, we proposed polyandry as a potential counterstrategy against t deleterious effects. Here, we show that (1) the efficiency of polyandry in reducing the t frequency strongly depends on the selective context and (2) polyandry helps to reduce male-biased leverage in sex dependent selection.

The decision where to live has far-reaching fitness consequences for animals. In contrast to most other mammals or birds that use sheltered nest sites, female Bechstein's bats frequently switch day roosts during one breeding season, and therefore must often decide where to spend the day. Selecting the right roost is important, because roost quality, e.g. microclimatic condition, influences survival and reproduction in bats. Although thermal factors are very important for the quality of roosts occupied by bats, whether bats base their day roost selection directly on roost temperature has not been tested in the field. Over one summer, we examined and tested the roost choice of 21 individually marked female Myotis bechsteinii living in one maternity colony. In a field experiment, we allowed the bats to choose between relatively warm versus cold bat boxes, while controlling for site preferences. We expected females to exhibit a preference for warm roosts during pregnancy and lactation to accelerate gestation and shorten the period of growth of their young. Roost occupancy over 160 census days reflected significant temperature differences among 89 surveyed roosts (14 tree holes and 75 bat boxes), and preferences changed with the season. Females significantly preferred cold roosts before parturition, whereas post-partum, they significantly favoured warm roosts. Temperature preferences were independent of the roost site, and thus roost selection was based directly on temperature. Boxes with significantly different daytime temperatures did not differ significantly at night. Consequently, bats would have to spend at least 1 day in a new roost to test it. Information transfer among colony members might facilitate knowledge of roost availability. Access to many roosts providing different microclimates is likely to be important for successful reproduction in the endangered Bechstein's bat.

In many studies with animals kept in groups, scientists need information about each individual's food access without disturbance or separation of the animals. We developed an automatic feeding device that allows measurement of individual food consumption and experimental manipulation of individual food availability in small social mammals, such as house mice. The feeding device is based on radio frequency identification that triggers access to a motor-driven metal arm filled with food pellets and is mediated with the help of subcutaneously implanted transponders.

According to theory in life-history and animal personality, individuals with high fitness expectations should be risk-averse, while individuals with low fitness expectations should be more bold. In female house mice, a selfish genetic element, the t haplotype, is associated with increased longevity under natural conditions, representing an appropriate case study to investigate this recent theory empirically. Following theory, females heterozygous for the t haplotype (+/t) are hypothesised to express more reactive personality traits and be more shy, less explorative and less active compared to the shorter-lived homozygous wildtype females (+/+). As males of different haplotype do not differ in survival, no similar pattern is expected. We tested these predictions by quantifying boldness, exploration, activity, and energetic intake in both +/t and +/+ mice. +/t females, unlike +/+ ones, expressed some reactive-like personality traits: +/t females were less active, less prone to form an exploratory routine and tended to ingest less food. Taken together these results suggest that differences in animal personality may contribute to the survival advantage observed in +/t females but fail to provide full empirical support for recent theory.

In house mice, genetic compatibility is influenced by the t haplotype, a driving selfish genetic element with a recessive lethal allele, imposing fundamental costs on mate choice decisions. Here, we evaluate the cost of genetic incompatibility and its implication for mate choice in a wild house mice population. In laboratory reared mice, we detected no fertility (number of embryos) or fecundity (ability to conceive) costs of the t, and yet we found a high cost of genetic incompatibility: heterozygote crosses produced 40% smaller birth litter sizes because of prenatal mortality. Surprisingly, transmission of t in crosses using +/t males was influenced by female genotype, consistent with postcopulatory female choice for + sperm in +/t females. Analysis of paternity patterns in a wild population of house mice showed that +/t females were more likely than +/+ females to have offspring sired by +/+ males, and unlike +/+ females, paternity of their offspring was not influenced by +/t male frequency, further supporting mate choice for genetic compatibility. As the major histocompatibility complex (MHC) is physically linked to the t, we investigated whether females could potentially use variation at the MHC to identify male genotype at the sperm or individual level. A unique MHC haplotype is linked to the t haplotype. This MHC haplotype could allow the recognition of t and enable pre- and postcopulatory mate choice for genetic compatibility. Alternatively, the MHC itself could be the target of mate choice for genetic compatibility. We predict that mate choice for genetic compatibility will be difficult to find in many systems, as only weak fertilization biases were found despite an exceptionally high cost of genetic incompatibility.

The ecological success of the house mouse as an invasive species is linked to its commensal interaction with humans, which has resulted in the species reaching and thriving in places outside its natural regions of origin. House mice live in a large variety of habitats and are able to cope with various environmental conditions. This ability is due to the species’ high reproductive potential, which has been interpreted as an adaptation to a colonizing life strategy. House mice typically live in social groups, and social relationships among group members are structured by cooperation and by the existence and resolution of conflicts. Flexibility in social and especially maternal behaviour allows females to reproduce and successfully wean offspring under variable resource conditions and to adjust parental investment to changing situations. Due to female social partner choice, communal nursing and a weaning strategy that produces many young of a minimal weaning weight, females are able to raise relatively large litters despite an energetically highly demanding lactation period and high juvenile mortality. Such traits may have also contributed to the fact that the laboratory mouse is one of the most successful model species in medicine and biology.

Infectious disease outbreaks can be devastating because of their sudden occurrence, as well as the complexity of monitoring and controlling them. Outbreaks in wildlife are even more challenging to observe and describe, especially when small animals or secretive species are involved. Modeling such infectious disease events is relevant to investigating their dynamics and is critical for decision makers to accomplish outbreak management. Tularemia, caused by the bacterium Francisella tularensis, is a potentially lethal zoonosis. Of the few animal outbreaks that have been reported in the literature, only those affecting zoo animals have been closely monitored. Here, we report the first estimation of the basic reproduction number R 0 of an outbreak in wildlife caused by F. tularensis using quantitative modeling based on a susceptible-infected-recovered framework. We applied that model to data collected during an extensive investigation of an outbreak of tularemia caused by F. tularensis subsp. holarctica (also designated as type B) in a closely monitored, free-roaming house mouse (Mus musculus domesticus) population in Switzerland. Based on our model and assumptions, the best estimated basic reproduction number R 0 of the current outbreak is 1.33. Our results suggest that tularemia can cause severe outbreaks in small rodents. We also concluded that the outbreak self-exhausted in approximately three months without administrating antibiotics.

In addition to sexual selection, selection resulting from social interactions in contexts other than mating can be a potent evolutionary force. Such social selection processes are facilitated whenever individual fitness varies as a result of any form of social interactions. The choice of social partners for communal care of young is such a situation in which interactants potentially experience fitness variance. In this study, we investigated the existence and impact of female social partner choice and the potential for social selection to occur in the cooperatively breeding wild house mouse, Mus domesticus. We analysed patterns of individual associations in groups of females, and compared the reproductive behaviour of females grouped with either a preferred or a nonpreferred social partner over an experimental life span of half a year, using spatial association as a measure of preference. We predicted low reproductive competition among preferred social partners and high competition, reflected in lower reproductive success, among nonpreferred. Our results showed that female house mice displayed nonrandom preferences, and that social partner choice yielded significant fitness benefits. Females in pairs with a preferred partner had a significantly higher probability to give birth and to establish an egalitarian, cooperative relationship, resulting in higher reproductive success than females in nonpreferred pairs. This suggests that interactions among females are subject to social selection processes, driving the evolution of female traits.

In order to understand why animals are social and how group members interact with each other it is important to know their relatedness. However, few studies have investigated the genealogy in complete social groups of free-living animals with low reproductive skew. This holds particularly true for bats. Although almost all bat species are social, their sociobiology is not well understood. Because they are volant, nocturnal and have a rather cryptic life-style, bats are difficult to observe in the wild. Furthermore females are generally gregarious making genetic parent-offspring assignment a challenging task. We used genetic markers in combination with knowledge about age and colony membership of individually marked bats to construct pedigrees in completely sampled maternity colonies of Bechstein's bats (Myotis bechsteinii). Despite considerable fluctuations in population size, no immigration occurred over 5 years in four colonies living in close proximity. Additionally, confrontation tests showed that females of one maternity colony were able to detect and attempted to prevent the intrusion of foreign females into a roost they occupy. Although colonies were absolutely closed, and 75% of the colony members lived together with close relatives (r≥=0.25), mean colony relatedness was nearly zero (0.02). Average relatedness therefore is a poor estimator for the potential of kin selection in Bechstein's bat colonies and may be misleading when attempting to understand the social structure of animals living in groups where many members breed. Based on our results we discuss the potential adaptive value of living in closed societies with low reproductive skew.

To explain coloniality in birds and bats, communal roosts have been hypothesized to serve as information centers where colony members transfer knowledge about their habitat. If information is exchanged about food, individuals with low hunting success benefit by following successful foragers to their profitable feeding sites. Colony members should therefore forage together regularly and move in pairs or groups among different feeding places. In 1996 and 1997, we used radio-telemetry to study the nightly habitat use of ten adult female Bechstein's bats (Myotis bechsteinii) living in one maternity colony. Over several nights, each bat revisited its own foraging area, although females regularly switched day-roosts at the same time. Most individual foraging areas showed no, or only little, overlap with each other. Distances between individual activity centers were negatively and significantly correlated with the degree of individual day-roost association. Genetic similarity among colony members, determined using seven nuclear microsatellites, was significantly positively correlated with the degree of overlap among individual areas. Five females were repeatedly radio-tracked at different seasons, months apart. Even between years, all bats maintained their individual hunting areas. Because females are very loyal to their individual foraging areas and these areas are typically substantial distances from each other, information transfer about feeding sites is unlikely to be the crucial factor promoting coloniality. Therefore, other benefits of sociality like cooperative breeding may cause communal roosting. Strong fidelity to individual foraging areas suggests the importance of familiarity with the local habitat, and profound site knowledge could be a crucial "resource", promoting female philopatry in Bechstein's bats.

We present a microgeographic analysis of mitochondrial DNA (mtDNA) in Bechstein's bats using three sources of control region sequence variability, including a novel mtDNA microsatellite, to assess individual relatedness both within and among 10 maternity colonies. Comparison of marker variability among 268 adult females revealed little genetic variability within each colony. However, most colonies were clearly distinguished by colony-specific mitochondrial haplotypes (total n = 28). Low intracolony variability and strong haplotype segregation among colonies, was reflected by an extraordinary high FST of 0.68, indicating a very low intercolony dispersal rate of approximately one female in five generations. Haplotype distribution among 18 solitary males showed that males frequently disperse between colony locations, indicating the absence of dispersal barriers. Bechstein's bat maternity colonies are thus closed groups that comprise 20-40 females probably belonging to only one or, at most, two matrilines. The genetic population structure of Bechstein's bats is in agreement with the hypothesis that females seek familiar and, at least, partially related cooperation partners for raising their young. Alternatively strong philopatry might reflect the importance of profound roost or habitat knowledge for successful reproduction in female Bechstein's bats.

Out of all the complex phenomena displayed in the behaviour of animal groups, many are thought to be emergent properties of rather simple decisions at the individual level. Some of these phenomena may also be explained by random processes only. Here we investigate to what extent the interaction dynamics of a population of wild house mice (Mus domesticus) in their natural environment can be explained by a simple stochastic model. We first introduce the notion of perceptual landscape, a novel tool used here to describe the utilisation of space by the mouse colony based on the sampling of individuals in discrete locations. We then implement the behavioural assumptions of the perceptual landscape in a multi-agent simulation to verify their accuracy in the reproduction of observed social patterns. We find that many high-level features – with the exception of territoriality – of our behavioural dataset can be accounted for at the population level through the use of this simplified representation. Our findings underline the potential importance of random factors in the apparent complexity of the mice's social structure. These results resonate in the general context of adaptive behaviour versus elementary environmental interactions., PLoS Computational Biology, 8 (11), ISSN:1553-734X, ISSN:1553-7358

Mongolian gerbils (Meriones unguiculatus) are a widely used model species in parasitology and epilepsy research. Under standard laboratory housing conditions gerbils develop stereotypic digging, a behavioural abnormality, which is caused by the lack of a burrow in the laboratory cage. Here we show that gerbils reared with access to an opaque artificial burrow (nest-box with access tube) develop significantly less stereotypic digging than those reared with transparent artificial burrows. Subadult gerbils also preferred opaque artificial burrows to transparent ones. Based on these findings we developed an artificial burrow system that could be integrated into laboratory Makrolon Type IV cages to prevent the development of stereotypic digging in gerbils by addressing their behavioural needs. Faecal cortisol levels were measured as a non-invasive method of comparing stress reactions in gerbils given access to the new integrated artificial burrow system or to an equivalent transparent burrow. Behavioural differences were observed between gerbils in the two housing conditions, but faecal cortisol levels were unaffected. We conclude that simple refinement of housing is effective in improving gerbil welfare in the laboratory.