The yellow-bellied marmot (Marmota flaviventris) is a social, ground-dwelling squirrel that lives either individually or in kin groups of from two to five adult females. Philopatry and daughter recruitment lead to the formation and persistence of matrilines at habitat sites. By using 37 years of demographic data for 12 habitat sites, we could determine long-term trends in the effects of group size on two measures of fitness, survivorship and net reproductive rate, which otherwise are obscured by annual fluctuations in these measures. Both size and number of matrilines varied among sites and survivorship and net reproductive rate varied among sites and among matriline sizes. The role of social organization was explored further by examining the effect of matriline size, averaged over all years and sites, on fitness. For both survivorship and net reproductive rate the relationship with matriline size was curvilinear. Fitness increased with the increase in matriline size and then decreased in the largest groups. Decreased fitness in matrilines of four or five was associated with agonistic behavior, a large number of 2-year-old females in the social group, and reproductive suppression. There is no evidence that females acted to increase their fitness by increasing indirect fitness; i.e., by assisting relatives, but attempted to increase direct fitness. Direct fitness increased when mortality and fission of large matrilines reduced group size and the surviving females increased reproduction.
Because sociality in mammals is potentially costly due to intraspecific competition, parasite transmission, or suppressed reproduction (1), sociality must enhance inclusive fitness directly through the production of offspring and possibly indirectly through the production of nondescendant relatives when compared with nonsocial conspecifics. However, in many societies, sociality is associated with reproductive skew; a few individuals do all or most of the breeding (2). Reproductive skew may be associated with a trade-off between reproduction and survival. For example, highly social species of marmots, such as Marmota olympus and M. vancouverensis, which do not reproduce before age 3, have higher survival at ages 1-4 than the less social M. flaviventris, which reproduces at age 2 (3-5). Increased group size is widely associated with reduced per capita reproductive success; e.g., increased social complexity among species of ground-dwelling sciurids is associated with decreased litter size and a smaller proportion of females breeding (6). Among cooperatively breeding species reproduction may be limited to a single female in a group (3, 7), and per capita reproductive success is lower in large coteries than in small coteries of black-tailed prairie dogs (8). In contrast, by using 37 years of field observations, we report that net reproductive rate (R o ), a direct measure of fitness (9), initially increases with increased group size in the yellow-bellied marmot (Marmota flaviventris), then decreases in the largest groups. Furthermore, ...