Carsten Schradin

INTRASPECIFIC VARIATION IN THE SPATIAL AND SOCIAL ORGANIZATION OF THE AFRICAN STRIPED MOUSE

Abstract Social flexibility, that is, the expression of different types of social systems within one species, has been reported in several mammalian taxa, including rodents. However, sociality in rodents has been studied mostly in captivity and the results are often regarded as laboratory artifacts. We present field data for 2 populations of the striped mouse (Rhabdomys pumilio), a diurnal muroid rodent from southern Africa. The 2 populations inhabit different environments and demonstrate differences in social organization. R. pumilio in the arid succulent karoo lives in social groups, comprising multiple adults of both sexes that share 1 nest and the same territory. Striped mice in the moist grasslands of South Africa are solitary. Females inhabit exclusive territories and territories of males overlap those of several females; association between the sexes appears to be restricted to mating. Home ranges of females in the grasslands were 6 times larger and home ranges of males were 10 times larger than their counterparts in the succulent karoo. Onset of reproductive activity occurred earlier and at a significantly lighter body weight in grasslands. In the succulent karoo, sexually mature offspring remain in their natal group without reproducing. We suggest that group living in the succulent karoo is in response to habitat saturation and the benefits of philopatry, whereas living alone in the grasslands may be a response to limiting resources, such as food.

The Striped Mouse (Rhabdomys pumilio) From the Succulent Karoo, South Africa: A Territorial Group-Living Solitary Forager With Communal Breeding and Helpers at the Nest.

The authors studied the striped mouse (Rhabdomys pumilio) in the semiarid succulent karoo of South Africa. Mice forage alone, but they live in groups that share a common nest. Groups consist of 1 to 4 breeding females, 1 to 2 breeding males, and their offspring of both sexes, which remain in their natal group even after reaching adulthood, participating in territorial defense and nest building without showing signs of reproductive activity. Interactions are typically amicable and take place inside or in front of the nest. In contrast, encounters with mice from other groups are aggressive. Group living in the succulent karoo is possibly due to ecological constraints imposed by habitat saturation because of a year-round stable food supply as well as associated benefits of philopatry.

Paternal Care in the Social and Diurnal Striped Mouse (Rhabdomys pumilio): Laboratory and Field Evidence

Paternal care is rare in mammals, occurring mainly in carnivores and neotropical primates, in which the difficulties of long generation time and large individuals lead to small sample sizes. Here, the authors show that paternal care can be easily studied in the four-striped mouse (Rhabdomys pumilio) because (a) captive males show all the patterns of parental care as do females, with the obvious exception of nursing; (b) in the field, wild males act amicably toward juveniles and retrieve pups experimentally presented to them; (c) the striped mouse facilitates experimental approaches in captivity because it has a short generation period and can be kept in large numbers; and (d) the striped mouse is diurnal, not only making observations in captivity easier but also enabling direct observations in the field.

Territorial defense in a group-living solitary forager: who, where, against whom?

Territoriality is of great significance for many species and a characteristic of most group-living animals. Territoriality is thought to lead to increased reproductive success by defending a particular area containing critical resources. I describe several factors that influence territorial aggression in free-ranging striped mice (Rhabdomys pumilio), a group-living solitary forager. I induced territorial aggression by attracting mice of different groups using bait either at territory boundaries or in front of nests. Striped mice are territorial and make decisions about whether or not to attack a mouse from another group based upon several factors: (1) the sex of the opponent: males are much more likely to attack strange males than strange females, whereas no sex specific aggression was observed in females; (2) the body size of the opponent: striped mice are much more likely to attack a strange mouse that is lighter than themselves; and (3) the location of encounters: striped mice are much more likely to attack strangers, even those significantly heavier than themselves, in front of the nest than at territory boundaries. These variations in territorial responses between different types of individuals may be due to the different ultimate consequences of territorial aggression for different animals.

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.

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

1. Social groups typically form due to delayed dispersal of adult offspring when no opportunities for independent breeding exist, or the costs of dispersal are higher than the costs of remaining philopatric. Ecological constraints are thought to be a main reason for group-living in animals. 2. Reproductive competition within groups can induce high costs of philopatry, and is thought to be a main reason for solitary living. 3. Experimental manipulation of reproductive competition is difficult. One solution is to compare sociality between periods with and without reproductive competition. 4. Here, we show empirically in a 8-year field study that striped mice (Rhabdomys pumilio) of both sexes were group-living during the breeding season when population density (PD) was high but solitary living when PD was low, supporting the ecological constraints model. 5. After the breeding season, in the absence of reproductive competition, the positive correlation between PD and percentage of group-living striped mice was absent. Almost all striped mice were group-living even under very low population densities. This supports the reproductive competition model. 6. Ambient temperature, food availability and predation pressure, did not influence sociality. 7. In captivity, the costs of reproductive competition in communal groups include female infanticide and aggression between females. 8. We conclude that group-living is favoured by constraints imposed through habitat saturation and by its benefits (improved thermoregulation by huddling, group-territoriality and predator avoidance), and that reproductive competition is a major force favouring solitary living in striped mice.

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.

Prolactin and paternal care: Comparison of three species of monogamous new world monkeys (Callicebus cupreus, Callithrix jacchus, and Callimico goeldii).

The authors explored whether prolactin is associated with paternal care in 3 monkey species: titi monkey (Callicebus cupreus), common marmoset (Callithrix jacchus), and Goeldis monkey (Callimico goeldii). They compared prolactin levels in fathers before and after infant birth as well as between fathers and nonfathers. C. cupreus fathers carry infants almost exclusively, have higher prolactin levels than nonfathers, but show no prolactin increase after infant birth. C. goeldii fathers carry infants only after 3 weeks, show an increase in prolactin levels during the precarrying period, but do not have higher levels than nonfathers. C jacchus fathers are the primary carriers, have higher prolactin levels than nonfathers, and show a trend for a prolactin increase after the birth of infants. In conclusion, species differences in the patterns of prolactin secretion were evident and reflect the different paternal roles.

Female home range size is regulated by resource distribution and intraspecific competition: a long-term field study

The size of an individuals home range is an important feature, influencing reproduction and survival, but it can vary considerably among both populations and individuals. The factors accounting for such variation are still poorly understood, and comprehensive long-term field studies considering various environmental factors that influence home range size are rare. We investigated the effects of seasonality, availability of food, cover, number of direct neighbours and the relative individual body mass on home range sizes in 125 adult female striped mice, Rhabdomys pumilio, in South Africa from 2004 to 2008. We used radiotelemetry to estimate home range sizes, trapping to determine the number of direct neighbours, and plant surveys in every home range to determine availability of food and cover. Home ranges were smaller when food quantity was high, many territorial neighbours were present, females had a relatively small body mass and during the nonbreeding season. We conclude that the availability of food resources and intraspecific competition are the main factors influencing home range size in female striped mice. Females enlarged their home ranges when territorial neighbours were few, and there was a significant positive correlation between home range size and quantity of food plants. This indicates that home range size might not reflect the minimal trade-off between access to resources that allow for a females survival and lowest cost for defending and foraging in that area. Instead, we propose a hypothesis for future research that female striped mice occupy areas several times larger than needed to improve their fitness by providing resources for future offspring.

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.