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Infanticide can be a major influence upon the social structure of species in which females maintain long-term associations with males. Previous studies have suggested that female mountain gorillas benefit from residing in multimale groups because infanticide occurs when one-male groups disintegrate after the dominant male dies. Here we measure the impact of infanticide on the reproductive success of female mountain gorillas, and we examine whether their dispersal patterns reflect a strategy to avoid infanticide. Using more than 40 years of data from up to 70% of the entire population, we found that only 1.7% of the infants that were born in the study had died from infanticide during group disintegrations. The rarity of such infanticide mainly reflects a low mortality rate of dominant males in one-male groups, and it does not dispel previous observations that infanticide occurs during group disintegrations. After including infanticide from causes other than group disintegrations, infanticide victims represented up to 5.5% of the offspring born during the study, and they accounted for up to 21% of infant mortality. The overall rates of infanticide were 2-3 times higher in one-male groups than multimale groups, but those differences were not statistically significant. Infant mortality, the length of interbirth intervals, and the age of first reproduction were not significantly different between one-male versus multimale groups, so we found no significant fitness benefits for females to prefer multimale groups. In addition, we found limited evidence that female dispersal patterns reflect a preference for multimale groups. If the strength of selection is modest for females to avoid group disintegrations, than any preference for multimale groups may be slow to evolve. Alternatively, variability in male strength might give some one-male groups a lower infanticide risk than some multimale groups, which could explain why both types of groups remain common.

As wildlife populations are declining, conservationists are under increasing pressure to measure the effectiveness of different management strategies. Conventional conservation measures such as law enforcement and community development projects are typically designed to minimize negative human influences upon a species and its ecosystem. In contrast, we define \"extreme\" conservation as efforts targeted to deliberately increase positive human influences, including veterinary care and close monitoring of individual animals. Here we compare the impact of both conservation approaches upon the population growth rate of the critically endangered Virunga mountain gorillas (Gorilla beringei beringei), which increased by 50% since their nadir in 1981, from approximately 250 to nearly 400 gorillas. Using demographic data from 1967-2008, we show an annual decline of 0.7%±0.059% for unhabituated gorillas that received intensive levels of conventional conservation approaches, versus an increase 4.1%±0.088% for habituated gorillas that also received extreme conservation measures. Each group of habituated gorillas is now continuously guarded by a separate team of field staff during daylight hours and receives veterinary treatment for snares, respiratory disease, and other life-threatening conditions. These results suggest that conventional conservation efforts prevented a severe decline of the overall population, but additional extreme measures were needed to achieve positive growth. Demographic stochasticity and socioecological factors had minimal impact on variability in the growth rates. Veterinary interventions could account for up to 40% of the difference in growth rates between habituated versus unhabituated gorillas, with the remaining difference likely arising from greater protection against poachers. Thus, by increasing protection and facilitating veterinary treatment, the daily monitoring of each habituated group contributed to most of the difference in growth rates. Our results argue for wider consideration of extreme measures and offer a startling view of the enormous resources that may be needed to conserve some endangered species.

Evaluating the factors influencing the patterns of female dispersal in mammals is critical to understanding its importance for male and female reproductive strategies and the evolution of social systems. In western lowland gorillas, females emigrate from their natal group (natal dispersal) but also disperse between groups multiple times in their life (secondary dispersal). This strategy is uncommon in primates and may reflect females seeking to reduce feeding competition or female choice for males that can provide protection against infanticide. In this study, we test how inbreeding avoidance, group size, and male competitive ability could influence female dispersal in western gorillas, using 25 natal and 285 secondary dispersal events collected over 20 years at Mbeli Bai. We found that while all females dispersed out of their natal group, presumably to avoid inbreeding, females also left their group before having their first offspring when the dominant male was not their father, pointing towards unconditional dispersal by nulliparous females. Regarding secondary dispersal, we found that females were more likely to stay with males that were at the beginning of their tenure than transfer and more likely to disperse away from (presumably old) males that were near the end of their tenure or closer to their death. Females were also more likely to leave larger groups for smaller ones suggesting a potential effect of within-group feeding competition or avoidance of outsider males or predators. This study highlights the intersexual conflict found in gorillas: while female choice for high-quality males may influence the formation of their relatively rare social system, smaller group size may be more important for females than previously thought, which runs counter to the males’interest of having a high number of females.

Long-term studies of population dynamics can provide insights into life history theory, population ecology, socioecology, conservation biology and wildlife management. Here we examine 25 years of population dynamics of western gorillas at Mbeli Bai, a swampy forest clearing in Nouabalé-Ndoki National Park, the Republic of Congo. The Mbeli population more than doubled from 101 to 226 gorillas during the study. After adjusting for a net influx of gorillas into the study population, the increase represents an inherent growth rate of 0.7% per year, with 95% confidence limits between -0.7% and 2.6%. The influx of gorillas mainly involved immigration of individuals into existing study groups (social dispersal), but it also included the appearance of a few previously unknown groups (locational dispersal). The average group size did not change significantly during the study, which is consistent with the possibility that western gorillas face socioecological constraints on group size, even when the population is increasing. We found no significant evidence of density dependence on female reproductive success or male mating competition. The distribution of gorillas among age/sex categories also did not change significantly, which suggests that the population had a stable age structure. Our results provide evidence of population stability or growth for some western gorillas (albeit within a small area). The results highlight the value of law enforcement, long-term monitoring, and protected areas; but they do not diminish the importance of improving conservation for this critically endangered species.

In group-living mammals, individual efforts to maximize reproductive success result in conflicts and compromises between the sexes. Females utilize counterstrategies to minimize the costs of sexual coercion by males, but few studies have examined the effect of such behaviors on female reproductive success. Secondary dispersal by females is rare among group-living mammals, but in western gorillas, it is believed to be a mate choice strategy to minimize infanticide risk and infant mortality. Previous research suggested that females choose males that are good protectors. However, how much female reproductive success varies depending on male competitive ability and whether female secondary dispersal leads to reproductive costs or benefits has not been examined. We used data on 100 females and 229 infants in 36 breeding groups from a 20-year long-term study of wild western lowland gorillas to investigate whether male tenure duration and female transfer rate had an effect on interbirth interval, female birth rates, and offspring mortality. We found that offspring mortality was higher near the end of males’ tenures, even after excluding potential infanticide when those males died, suggesting that females suffer a reproductive cost by being with males nearing the end of their tenures. Females experience a delay in breeding when they dispersed, having a notable effect on birth rates of surviving offspring per female if females transfer multiple times in their lives. This study exemplifies that female counterstrategies to mitigate the effects of male-male competition and sexual coercion may not be sufficient to overcome the negative consequences of male behavior.

Socioecological models predict that contest competition for clumped foods can lead to higher energy intake and lower energy expenditure for higher-ranking individuals. Here, we examine the relationships between dominance rank and energy intake and expenditure of female mountain gorillas in Bwindi Impenetrable National Park, Uganda (Gorilla beringei beringei). Bwindi gorillas have weak dominance relationships, feed on nonreproductive plant parts throughout the year, and consume fruit when it is seasonally available. We used behavioral observations on one group of gorillas and nutritional analysis of their major food items to calculate energy intake rates and estimated energy expenditure. Using linear mixed models, we found a significant positive relationship between dominance rank and energy intake rates, due to higher-ranking females having faster ingestion rates, rather than consuming foods with higher energy concentrations. Lower-ranking females did not spend significantly more time feeding to compensate for their lower energy intake rates. Lower-ranking females spent significantly more time traveling than higher-ranking females, leading to a negative relationship between dominance rank and energy expenditure. The combined results revealed a significant positive relationship between dominance rank and energy balance. Higher-ranking females did not spend longer feeding on fruit than lower-ranking ones, and the relationship between dominance rank and energy intake rates was not stronger when fruit was available. According to socioecological models, these results suggest that contest competition may be occurring with both fruit and nonreproductive plant parts, which would be consistent with growing evidence that nonreproductive plant parts can be contestable.

Dispersal is a key event in the life of an animal and it influences individual reproductive success. Male mountain gorillas exhibit both philopatry and dispersal, resulting in a mixed one-male and multimale social organization. However, little is known about the relationship between male dispersal or philopatry and reproductive careers in Bwindi mountain gorillas. Here we analyze data spanning from 1993 to 2017 on social groups in Bwindi Impenetrable National Park, Uganda to examine the proportion of males that disperse, age of dispersal, pathways to attaining alpha status, fate of dispersing males and philopatric males, and male tenure length as well as make comparisons of these variables to the Virunga mountain gorilla population. We report previously undocumented cases of dispersal by immature males and old males and we also observed the only known case of a fully mature male immigrating into a breeding group. We used genetic tracking of known individuals to estimate that a minimum of 25% of males that disperse to become solitary males eventually form new groups. No differences were found between the Bwindi and Virunga population in the age of male dispersal, the proportion of males that disperse, the age of alpha male acquisition, and dominance tenure length. The lack of differences may be due to small sample sizes or because the observed ecological variability does not lead to life history differences between the populations. Males in both populations follow variable strategies to attain alpha status leading to the variable one-male and multimale social organization, including dispersal to become solitary and eventually form a group, via group fissioning, usurping another alpha male, or inheriting the alpha position when a previous group leader dies.

Sexual coercion and courtship are possible explanations for why male primates may direct agonistic behavior towards females. If so, then in species where females exhibit mate choice by transferring between males: (a) females who are not lactating (potential migrants) should receive more agonistic behavior than other females, (b) males should exhibit more agonistic behavior towards females during intergroup encounters than when no rival males are nearby, and (c) males should show more herding behavior when their group contains potential migrant females. We tested those hypotheses in a population of approximately 150 western gorillas at Mbeli Bai, northern Congo. We also tested whether difference in male phenotypic traits influenced their rates of agonistic behavior towards females. Of the 332 observed cases of male agonistic behavior towards females, 29 % represented feeding competition, 7 % involved interventions in conflicts between females, and the remaining 64 % were considered potential evidence of sexual coercion and/or courtship. After excluding the cases of feeding competition and intervention, a multivariate analysis indicated that potential migrant females received agonistic behavior at a statistically significantly higher rate than other adult females. Females also received agonistic behavior at a significantly higher rate during intergroup encounters than at other times. Herding occurred during 22 % of the 292 dyadic interunit encounters, and was significantly more likely to occur when the group contained a potential migrant female, but was not influenced by the number of adult females or the type of group encountered. Males with shorter body lengths had significantly higher rates of aggression, but phenotypic traits were not significantly correlated with herding. Collectively, our results are consistent with sexual coercion and/or courtship as an explanation for male-to-female agonistic behavior, but we are unable to distinguish between those two male mating strategies. Both types of behavior are likely due to a combination of perceived risks of female transfer and opportunity to advertise protector ability.

Using 30 years of demographic data from 15 groups, this study estimates how harem size, female fertility, and offspring survival may contribute to variance in the siring rates of dominant male mountain gorillas throughout the Virunga Volcano Region. As predicted for polygynous species, differences in harem size were the greatest source of variance in the siring rate, whereas differences in female fertility and offspring survival were relatively minor. Harem size was positively correlated with offspring survival, even after removing all known and suspected cases of infanticide, so the correlation does not seem to reflect differences in the ability of males to protect their offspring. Harem size was not significantly correlated with female fertility, which is consistent with the hypothesis that mountain gorillas have minimal feeding competition. Harem size, offspring survival, and siring rates were not significantly correlated with the proportion of dominant tenures that occurred in multimale groups versus one-male groups; even though infanticide is less likely when those tenures end in multimale groups than one-male groups. In contrast with the relatively small contribution of offspring survival to variance in the siring rates of this study, offspring survival is a major source of variance in the male reproductive success of western gorillas, which have greater predation risks and significantly higher rates of infanticide. If differences in offspring protection are less important among male mountain gorillas than western gorillas, then the relative importance of other factors may be greater for mountain gorillas. Thus, our study illustrates how variance in male reproductive success and its components can differ between closely related species.

The effect of feeding competition on foraging efficiency is an important link between ecological factors and the social organization of gregarious species. We examined the effects of group size on daily travel distances, activity budgets, and energy intake of mountain gorillas in Rwanda. We measured daily travel distances of five groups, activity budgets of 79 gorillas in nine groups, and energy intake data for 23 adult females in three groups over a 16-month period. Travel distances and the proportion of time spent traveling increased with size for most groups, which would be expected if their foraging efficiency is limited by intragroup feeding competition. However, travel distances and times decreased for the largest group, which also had higher energy intake rates than intermediate sized groups. The improved foraging efficiency of the largest group may be explained by advantages in intergroup contest competition. The largest group had much lower home range overlap than the other study groups which may be due to groups avoiding one another as a result of male mating competition. Collectively, our results indicate that intermediate sized groups had the lowest foraging efficiency and provide a new twist on the growing evidence of non-linear relationships between group size and foraging efficiency in primates.