Why Class Formation Occurs in Humans but Not among Other Primates : A Primate Coalitions Model.

Most human societies exhibit a distinct class structure, with an elite, middle classes, and a bottom class, whereas animals form simple dominance hierarchies in which individuals with higher fighting ability do not appear to form coalitions to "oppress" weaker individuals. Here, we extend our model of primate coalitions and find that a division into a bottom class and an upper class is inevitable whenever fitness-enhancing resources, such as food or real estate, are exploitable or tradable and the members of the bottom class cannot easily leave the group. The model predicts that the bottom class has a near flat, low payoff and always comprises at least half the society. The upper class may subdivide into one or more middle class(es), resulting in improved payoff for the topmost members (elite). The model predicts that the bottom class on its own is incapable of mounting effective counter-coalitions against the upper class, except when receiving support from dissatisfied members of the middle class(es). Such counter-coalitions can be prevented by keeping the payoff to the lowest-ranked members of the middle classes (through concessions) well above that of the bottom class. This simple model explains why classes are also absent in nomadic hunter-gatherers and predominate in (though are not limited to) societies that produce and store food. Its results also agree well with various other known features of societies with classes.

The Conditions Favoring Between-Community Raiding in Chimpanzees, Bonobos, and Human Foragers.

Chimpanzees, bonobos, and human foragers share a fission-fusion social system and a mating system of joint male resource defense polygyny. Within-community skew in male strength varies among and within species. In this study, we extend a mathematical model of within-group male coalition formation among primates to derive the conditions for between-community conflicts in the form of raids. We show that the main factor affecting the presence of successful raiding is the likelihood of major discrepancies in party strength, which are set by party size distributions (and thus community size) and the skew in strength. This study confirms the functional similarities between the raiding of chimpanzees and human foragers, and it supports the "imbalance of power" hypothesis for raiding. However, it also proposes two amendments to this model. First, the absence of raiding in bonobos may be attributable more to potential female involvement in defense against raids, which increases the size of defensive coalitions. Second, the model attributes some of the raiding in humans to major contrasts in instantaneous fighting ability created by surprise raids on unarmed victims; it also draws attention to the distinction between minor raids and major raids that involve multiple bands of the same community.

Why do chimpanzee males attack the females of neighboring communities?

Our closest nonhuman primate relatives, chimpanzees, engage in potentially lethal between-group conflict; this collective aggressive behavior shows parallels with human warfare. In some communities, chimpanzee males also severely attack and even kill females of the neighboring groups. This is surprising given their system of resource defense polygyny, where males are expected to acquire potential mates. We develop a simple mathematical model based on reproductive skew among primate males to solve this puzzle. The model predicts that it is advantageous for high-ranking males but not for low-ranking males to attack females. It also predicts that more males gain a benefit from attacking females as the community's reproductive skew decreases, i.e., as mating success is more evenly distributed. Thus, fatal attacks on females should be concentrated in communities with low reproductive skew. These attacks should also concur with between-community infanticide. A review of the chimpanzee literature provides enough preliminary support for this prediction to warrant more detailed testing.

Social organization and the evolution of cumulative technology in apes and hominins.

Culturally supported accumulation (or ratcheting) of technological complexity is widely seen as characterizing hominin technology relative to that of the extant great apes, and thus as representing a threshold in cultural evolution. To explain this divide, we modeled the process of cultural accumulation of technology, which we defined as adding new actions to existing ones to create new functional combinations, based on a model for great ape tool use. The model shows that intraspecific and interspecific variation in the presence of simple and cumulative technology among extant orangutans and chimpanzees is largely due to variation in sociability, and hence opportunities for social learning. The model also suggests that the adoption of extensive allomaternal care (cooperative breeding) in early Pleistocene Homo, which led to an increase in sociability and to teaching, and hence increased efficiency of social learning, was enough to facilitate technological ratcheting. Hence, socioecological changes, rather than advances in cognitive abilities, can account for the cumulative cultural changes seen until the origin of the Acheulean. The consequent increase in the reliance on technology could have served as the pacemaker for increased cognitive abilities. Our results also suggest that a more important watershed in cultural evolution was the rise of donated culture (technology or concepts), in which technology or concepts was transferred to naïve individuals, allowing them to skip many learning steps, and specialization arose, which allowed individuals to learn only a subset of the population's skills.

A model for the evolution of developmental arrest in male orangutans.

Male Sumatran orangutans (Pongo abelii) may delay for many years the acquisition of the full array of secondary sexual traits, including their characteristic cheek flanges. Such flexible developmental arrest is unique among male primates. Among male Bornean orangutans (Pongo pygmaeus) such long delays appear less common. Here, we develop a simple model to identify the conditions under which developmental arrest can be adaptive. We show that the baseline strategy (i.e., males are not susceptible to arrest) cannot be invaded by the flexible strategy (i.e., males can arrest their development when the conditions are unfavorable) when the potential for high-ranking unflanged or flanged males to monopolize sexual access to females is low. In contrast, at high monopolization potential, the flexible strategy is the evolutionarily stable strategy. We also derive the proportion of flanged males in the population for each combination of monopolization values. This model concurs with field data that found a different monopolization potential between Bornean and Sumatran flanged males and a lower proportion of flanged males in the population in Sumatran orangutans. Pronounced developmental arrest is linked to very low adult mortality, which explains why it is so limited in its taxonomic distribution.

Mating strategies in primates: a game theoretical approach to infanticide.

Infanticide by newly immigrated or newly dominant males is reported among a variety of taxa, such as birds, rodents, carnivores and primates. Here we present a game theoretical model to explain the presence and prevalence of infanticide in primate groups. We have formulated a three-player game involving two males and one female and show that the strategies of infanticide on the males' part and polyandrous mating on the females' part emerge as Nash equilibria that are stable under certain conditions. Moreover, we have identified all the Nash equilibria of the game and arranged them in a novel hierarchical scheme. Only in the subspace spanned by the males are the Nash equilibria found to be strict, and hence evolutionarily stable. We have therefore proposed a selection mechanism informed by adaptive dynamics to permit the females to transition to, and remain in, optimal equilibria after successive generations. Our model concludes that polyandrous mating by females is an optimal strategy for the females that minimizes infanticide and that infanticide confers advantage to the males only in certain regions of parameter space. We have shown that infanticide occurs during turbulent changes accompanying male immigration into the group. For changes in the dominance hierarchy within the group, we have shown that infanticide occurs only in primate groups where the chance for the killer to sire the next infant is high. These conclusions are confirmed by observations in the wild. This model thus has enabled us to pinpoint the fundamental processes behind the reproductive decisions of the players involved, which was not possible using earlier theoretical studies.

A model for tool-use traditions in primates: implications for the coevolution of culture and cognition.

Inspired by the demonstration that tool-use variants among wild chimpanzees and orangutans qualify as traditions (or cultures), we developed a formal model to predict the incidence of these acquired specializations among wild primates and to examine the evolution of their underlying abilities. We assumed that the acquisition of the skill by an individual in a social unit is crucially controlled by three main factors, namely probability of innovation, probability of socially biased learning, and the prevailing social conditions (sociability, or number of potential experts at close proximity). The model reconfirms the restriction of customary tool use in wild primates to the most intelligent radiation, great apes; the greater incidence of tool use in more sociable populations of orangutans and chimpanzees; and tendencies toward tool manufacture among the most sociable monkeys. However, it also indicates that sociable gregariousness is far more likely to produce the maintenance of invented skills in a population than solitary life, where the mother is the only accessible expert. We therefore used the model to explore the evolution of the three key parameters. The most likely evolutionary scenario is that where complex skills contribute to fitness, sociability and/or the capacity for socially biased learning increase, whereas innovative abilities (i.e., intelligence) follow indirectly. We suggest that the evolution of high intelligence will often be a byproduct of selection on abilities for socially biased learning that are needed to acquire important skills, and hence that high intelligence should be most common in sociable rather than solitary organisms. Evidence for increased sociability during hominin evolution is consistent with this new hypothesis.

Shape anisotropy of lipid molecules and voids.

Biological polymers, viz., proteins, membranes and micelles exhibit structural discontinuities in terms of spaces unfilled by the polymeric phase, termed voids. These voids exhibit dynamics and lead to interesting properties which are experimentally demonstrable. In the specific case of phospholipid membranes, numerical simulations on a two-dimensional model system showed that voids are induced primarily due to the shape anisotropy in binary mixtures of interacting disks. The results offer a minimal description required to explain the unusually large permeation seen in liposomes made up of specific lipid mixtures (Mathai & Sitaramam, 1994). The results are of wider interest, voids being ubiquitous in biopolymers.

Mode locking of spatiotemporally periodic orbits in coupled sine circle map lattices.

We study the organization of mode-locked intervals corresponding to the stable spatiotemporally periodic solutions in a lattice of diffusively coupled sine circle maps with periodic boundary conditions. Spatially periodic initial conditions settle down to spatiotemporally periodic solutions over large regions of the parameter space. In the case of synchronized solutions resulting from synchronized initial conditions, the mode-locked intervals have been seen to follow strict Farey ordering in the temporal periods. However, the nature of the organization of the mode-locked intervals corresponding to higher spatiotemporal periods is highly dependent on initial conditions and on system parameters. Farey ordering in the temporal periods is seen at low coupling for mode-locked intervals of all spatial periods. On the other hand, stable spatial period two solutions show an interesting reversal of Farey ordering at high values of coupling. Other spatially periodic solutions show a complete departure from Farey ordering at high coupling. We also examine the issue of completeness of the mode-locked intervals via a calculation of the fractal dimension of the complement of the mode-locked intervals as a function of the coupling epsilon and the nonlinearity parameter K. Our results are consistent with completeness over a range of values for these parameters. Spatiotemporally periodic solutions of the traveling wave type have their own organization in the parameter space. Novel bifurcations to other types of solutions are seen in the mode-locked intervals. We discuss various features of these bifurcations. We also define a set of new variables using which an analytic treatment of the bifurcations along the Omega=0 line is carried out.