Research on lions by Heinsohn and Packer (1995) illustrates the behavioral complexities displayed by animals that are simultaneously territorial and cooperative group-foragers. African lions (Panthera leo) engage in a wide variety of group-level activities from hunting to communal cub rearing. At the same time, the group defends a territory from other groups of lions. When prey is difficult to capture they hunt cooperatively, but cooperation breaks down when prey is easy to catch. Female lions nurse each other's young, but, more importantly, they jointly protect their young from males that are intent upon infanticide. The threat of attack by conspecifics is a driving force in lion sociality. Large prides dominate smaller ones, and solitary lions are often killed or injured during attacks by lions of the same sex.
Using playbacks of recorded roars, Heinsohn and Packer found that lions are able to distinguish pride members from strangers. They also found that certain females show a consistent behavior of lagging behind their companions during group activities, including hunting.
Female lions live in social groups (prides), which contain 3-6 related adults, their dependent offspring, and a group of immigrant males. The males defend the pride against incursions by other males; females defend their young against infanticidal males, and the territory from other females. At least two females are needed for a territory, and they advertise ownership by roaring. Using broadcasted roars, the investigators showed that some females become "laggards" early in life, and this behavior persists into adulthood. Laggards were those individuals that hung back, and approached the audio speaker only after the leaders had already responded. The order in which the individuals approached the speaker was the same throughout the playbacks. Because territorial fights often lead to injury or death, laggards were ensuring their safety, at least from initial attacks. They typically followed the leaders by 30 to 120 seconds.
In the theoretical game or model known as "prisoner's dilemma," in any single task two individuals benefit when they work together (mutual cooperation) but both lose when neither contributes (mutual defection). However, in this game, the greatest payoff for one individual comes from providing no help (cheating) to a partner who cooperates, while the lowest payoff results from helping out (cooperating) while the partner cheats. In a repeated series of encounters, however, cheaters are eventually punished by withdrawal of further cooperation by other individuals. In large groups the game gets more complicated, but cheaters can eventually be detected and punished.
In the case of the lion pride, female leaders approached the speaker slowly and stopped to look behind at the laggards. The leader females mistrust the laggards, but they do not directly punish them. Although lions might be tempted to reduce their risk of injury from territorial defense by withdrawing or not cooperating, they still need companions for territorial defense, to share the protection of the young from males, and for the capture of large prey. In addition, females are often closely related, usually sisters. Given their shared genes, laggards appear to be tolerated, and leaders continue to arrive alone at the speaker 30 seconds to two minutes before the laggards arrive.
Heinsohn and Packer classified the behavior of individual female lions as: (i) unconditional cooperators, who always led the response; (ii) unconditional laggards, who always lagged behind; (iii) conditional cooperators, who lagged least when needed most; (iv) conditional laggards, who lagged most when needed most.
The parallels with human societies are evident. How these behaviors developed and why they are tolerated in animal (as well as human!) societies will remain a fascinating topic for future investigations.
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