Better off alone! Reproductive competition and ecological constraints determine sociality in the African striped mouse (Rhabdomys pumilio).

1. While the reasons for group-living have been studied for decades, little is known about why individuals become solitary. 2. Several previous experimental studies could demonstrate that group-living can arises as a consequence of ecological constraints. 3. It has been argued that reproductive competition between group members leads to significant costs of group-living, being a main reason of solitary-living. However, so far, no studies tested experimentally whether reproductive competition can explain solitary-living. 4. Using a socially flexible species, the African striped mouse (Rhabdomys pumilio), we tested experimentally in the field whether dispersal and solitary-living are more likely to occur when reproductive competition is present. 5. We investigated ecological constraints, here expressed as a function of population density, by removing groups of striped mice and creating vacant territories. To control for the effect of reproductive competition, which occurs only during the breeding season, we performed experiments during both the breeding and the non-breeding season. This is the first removal experiment performed in a species with communal breeding during the non-breeding season. 6. During the breeding season, when population density was low, more striped mice from experimental groups moved into the vacant territories and became solitary than striped mice from control groups. This is in support of the ecological constraints hypothesis. 7. During the non-breeding season, striped mice remained group-living despite the availability of free territories. Significantly, more striped mice became solitary-living during the breeding than during the non-breeding season. This is the first experimental support for the reproductive competition hypothesis explaining solitary-living. 8. Analysis of the sexual maturity of males showed that males which became solitary had a higher reproductive potential than males that remained group-living. Analysis of the body mass data of females showed that more solitary females reproduced than group-living females. These results indicate that by becoming solitary individuals of both sexes avoided costs of reproductive competition within groups. 9. Our study provides experimental evidence that reproductive competition within groups can lead to dispersal and solitary-living.

Schoepf I ，Schradin C 《-》

Reproductive competition favours solitary living while ecological constraints impose group-living in African striped mice.

Schradin C ，König B ，Pillay N 《-》

Endocrinology of sociality: comparisons between sociable and solitary individuals within the same population of African striped mice.

The social organization of species ranges from solitary-living to complex social groups. While the evolutionary reasons of group-living are well studied, the physiological mechanisms underlying alternative social systems are poorly understood. By studying group-living and solitary individuals of the same species, we can determine hormonal correlates of sociality without the problem of confounding phylogenetic factors. The African striped mouse (Rhabdomys pumilio) is a socially flexible species, which can be solitary or alternatively form complex family groups, depending on population density and the extent of reproductive competition. We predicted group-living striped mice to show signs of reproductive suppression and social stress, resulting in higher corticosterone but lower testosterone levels when compared to solitary-living individuals. To determine whether differences in social organization correlated with hormonal differences, we collected blood samples from free-living striped mice during four breeding seasons when we experimentally induced solitary-living in philopatric individuals by locally reducing population density. Striped mice that were group-living did not change their corticosterone or estosterone levels during the study, indicating that there was no temporal effect during the breeding season. Striped mice of both sexes had significantly lower corticosterone levels after switching from group- to solitary-living. Solitary males - but not solitary females - had higher testosterone levels than group-living conspecifics. Our results suggest that group-living results in physiological stress and can induce reproductive suppression, at least in philopatric males. The switch to solitary-living may thus be a tactic to avoid reproductive competition within groups, and is associated with decreased stress hormone levels and onset of independent reproduction.

Schoepf I ，Schradin C 《-》

Alternative reproductive tactics in female striped mice: heavier females are more likely to breed solitarily than communally.

Alternative reproductive tactics (ARTs) are discrete reproductive phenotypes governed by decision rules called strategies. ARTs are fixed for life in species with alternative strategies, while tactic expression is plastic in species with a single strategy. ARTs have been investigated in males of many species, but few studies have tested whether the same theoretical framework applies in females. Female striped mice (Rhabdomys pumilio) employ three ARTs: communal breeders give birth in a nest shared with female kin and a breeding male and show allo-parental care; returners give birth away from the shared nest and later return to it; and solitary breeders give birth away from the shared nest and do not return to the group. Here, studying free-living female striped mice over six breeding seasons, we tested whether ARTs arise from alternative strategies or a single strategy. We also asked to what extent stochastic extrinsic factors explain whether individuals become solitary rather than group living. Females switched tactics, consistent with a single strategy, so we tested whether this represented a mixed or conditional single strategy. Only the latter predicts differences between ARTs in traits indicating competitive ability, such as body mass or age, before individuals adopt a tactic. We weighed females at conception when they were still group living to eliminate potential confounding effects of gestation and subsequent social tactic (solitary versus group living) on body mass. Females that went on to use a solitary ART were heavier than those that became communal breeders and returners, in support of a conditional strategy. Solitary breeders also arose through extrinsic factors (mortality of all adult female group members). They weighed less than females that became solitary while relatives were alive, but did not differ in body mass from communal breeders and returners. We conclude that ART theory applies to both sexes, with female striped mice following a conditional single strategy. Future studies should consider the possibility that phenotypes that superficially resemble evolved tactics might also arise through non-adaptive extrinsic causes.

Hill DL ，Pillay N ，Schradin C 《-》

Social flexibility and social evolution in mammals: a case study of the African striped mouse (Rhabdomys pumilio).

Environmental change poses challenges to many organisms. The resilience of a species to such change depends on its ability to respond adaptively. Social flexibility is such an adaptive response, whereby individuals of both sexes change their reproductive tactics facultatively in response to fluctuating environmental conditions, leading to changes in the social system. Social flexibility focuses on individual flexibility, and provides a unique opportunity to study both the ultimate and proximate causes of sociality by comparing between solitary and group-living individuals of the same population: why do animals form groups and how is group-living regulated by the environment and the neuro-endocrine system? These key questions have been studied for the past ten years in the striped mouse Rhabdomys pumilio. High population density favours philopatry and group-living, while reproductive competition favours dispersal and solitary-living. Studies of genetic parentage reveal that relative fitness of alternative reproductive tactics depends on the prevailing environment. Tactics have different fitness under constrained ecological conditions, when competitive ability is important. Under conditions with relaxed ecological constraints, alternative tactics can yield equal fitness. Both male and female striped mice display alternative reproductive tactics based on a single strategy, i.e. all individuals follow the same decision rules. These changes are regulated by endocrine mechanisms. Social flexibility is regarded as an adaptation to unpredictably changing environments, selecting for high phenotypic flexibility based on a broad reaction norm, not on genetic polymorphism for specific tactics.

Schradin C ，Lindholm AK ，Johannesen J ，Schoepf I ，Yuen CH ，König B ，Pillay N ... -  《-》