Abstract
This chapter is the first of five comprising Part III. Though, as in Chap. 6, we have previously allowed some lexical analyses to interpolate Part II’s historical-empirical thrust, Part III is predominately statistical, even as it continues to review relevant literature and history. Though, consistent with the mandate of this monograph, we aim ultimately to establish the reality of human group selection, this initial chapter alone treats chimpanzees. To thoroughgoing evolutionists, the relevance will be self-evident; we only add that establishing evidence of group selection in such a highly related species foundationally supports the empirical argument for human group selection, as presented in the four subsequent chapters constituting Part III of this volume.
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Notes
- 1.
Chimpanzees inhabit multimale-multifemale communities characterized for their fission-fusion dynamics, within which individuals decrease the intensity of intragroup competition by foraging or exploring in subgroups (Aureli et al., 2008; Lehmann & Boesch, 2004; Lehmann, Korstjens, & Dunbar, 2007). Chimpanzees are polygynandrous, with males and females copulating with multiple individuals (Van Schaik, 2016). Although male reproductive skewness has been reported, males also employ an array of alternative mating tactics such as sperm competition (Dixson, 2012; Muller & Pilbeam, 2017) and collective mate guarding (Watts, 1998). In contrast to other primates living in polygynous societies, where a single male has a reproductive monopoly, chimpanzees display an attenuation in their sexual dimorphism (e.g., in canine size; Plavcan, 2001, 2012; Plavcan, Van Schaik, & Kappeler, 1995). Chimpanzees exhibit sex-biased dispersal, with females abandoning their natal group after reaching sexual maturity (Langergraber, Mitani, & Vigilant, 2009; Mitani, Watts, & Muller, 2002; Pusey, 1980). Male philopatry has considerable social sequelae such as the development of intracommunity coalitions and alliances (Chapais, 2009; Gilby et al., 2013; Wilson & Glowacki, 2017).
- 2.
- 3.
The annexation of this area generated several behavioral changes. For example, Ngogo chimpanzees spent over 30% of the observation time foraging and socializing in the captured region, a pattern that lasted for at least five months (Mitani et al., 2010).
- 4.
Julian, Badfoot, and Light Brown, in 1991, 1998, and 2001
- 5.
A similar approach was adopted in previous publications. For example, Wrangham et al. (2006) generated a comprehensive cross-site database describing instances of intracommunity and intercommunity lethal aggression from five sites and nine communities. Following traditional epidemiological procedures, the authors estimated the sites had a median mortality rate of 69 per 100,000 per year, based on observed and inferred cases, and 287 per 100,000 per year, including suspected cases (Wrangham et al., 2006). Communities also exhibited noticeable differences in killing rates. Even though the total rate across communities ranged from 125 to 306 per 100,000 per year, some communities experienced higher rates than others (Wrangham et al., 2006). Kahama, for example, reached a value of 12,000 per 100,000 per year, while Sonso did not experience any attack (Wrangham et al., 2006). Across communities, adult and adolescent males were often the victims of these attacks, with a rate of 355 per 100,000 per year, followed by infants and juveniles with 92, and adult and adolescent females with 28. This value stands in contrast to the frequency of intracommunity lethal aggression, with infants and juveniles displaying a median rate of 429 per 100,000 per year, as compared to adult and adolescent males with 254 per 100,000 per year (Wrangham et al., 2006).
- 6.
Though this pattern generalizes to other chimpanzee communities, regional differences exist between Eastern and Western communities. In contrast to chimpanzees from Gombe, female chimpanzees at the Taï National Park experience less severe, life-threatening attacks (Boesch et al., 2008). Moreover, the frequency of sexual interactions between neighboring communities at Taï is five times greater than that of Eastern communities (Boesch et al., 2008). These behavioral differences could be attributed to socioecological variations. Taï chimpanzees forage in larger parties and exhibit more social cohesion (Boesch, 1991; Boesch et al., 2008). Higher levels of gregariousness allow vulnerable individuals to be rescued by nearby supporters during intercommunity encounters, a phenomenon uncommon in Eastern communities (Boesch et al., 2008). Taï’s larger group size could also be attributed to higher predation rates (Boesch, 1991). It is worth noting, however, that even under circumstances of greater social cohesion, intercommunity killings do occur (Boesch et al., 2007, 2008).
- 7.
Lanchester’s “linear law” predicts that the largest group will not deploy all its units in a battle. Victory will depend on the relative difference in fighting force between the factions (Wilson, Britton, & Franks, 2002). According to the “square law,” if one of the groups outnumbers the other, the largest group should allocate all its units in a concentrated attack. The numerical advantage will influence the outcome of the conflict (Wilson et al., 2002).
- 8.
Given that previous publications have used reproductive success as a proxy for fitness in chimpanzees (Gilby et al., 2013), the present study employed a similar approach by using the males’ paternity success as a surrogate for the individual’s fitness.
- 9.
We would like to thank Robyn Stea for help in coding these data and JohnMichael Jurgensen for his feedback on this chapter.
- 10.
Even though traditional factor analyses rely on the extraction of latent variables from observable indicators across individuals (an R-type matrix), it is also statistically feasible to determine underlying groups by examining the correlations between individuals across occasions (an S-type matrix; Gorsuch, 2015).
- 11.
Langergraber et al. (2017) squared this variable. The present chapter retained this transformation.
- 12.
Researchers have also argued that the persistence of lethal intercommunity competition arises from the low costs accrued by raiding males when targeting vulnerable or solitary individuals in the rival group. Even though the experimental and observational evidence endorses perspectives concentrating on examining the low fitness costs of ambushes and incursions, such as the imbalance of power hypothesis, these results should not discourage researchers from further examining the fitness benefits obtained by raiding males. Similarly, future studies should consider the role of multilevel selection in the evolution of chimpanzee intercommunity competition.
- 13.
See Sober and Wilson (1998), for a detailed overview of the logic inconsistencies associated with the parsimony argument.
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Peñaherrera-Aguirre, M., Figueredo, A.J., Hertler, S.C. (2020). Chimpanzee Intercommunity Conflict: Fitness Outcomes, Power Imbalances, and Multilevel Selection. In: Multilevel Selection. Palgrave Macmillan, Cham. https://doi.org/10.1007/978-3-030-49520-6_8
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