Social evolution leads to persistent corruption.

Cooperation can be sustained by institutions that punish free-riders. Such institutions, however, tend to be subverted by corruption if they are not closely watched. Monitoring can uphold the enforcement of binding agreements ensuring cooperation, but this usually comes at a price. The temptation to skip monitoring and take the institution's integrity for granted leads to outbreaks of corruption and the breakdown of cooperation. We model the corresponding mechanism by means of evolutionary game theory, using analytical methods and numerical simulations, and find that it leads to sustained or damped oscillations. The results confirm the view that corruption is endemic and transparency a major factor in reducing it.

How Virtue Was Born.

This paper summarizes the Opening Lecture of the European Forum Alpbach 2017 in Tyrol/Austria (https://www.alpbach.org/de/). It deals with the evolution of cooperation throughout the history of life on Earth, and in particular human cooperation based on partnership. It emphasizes the role of institutions providing incentives for cooperation, and the role of praise and blame in guiding our actions. This helps for a better understanding of the social contract, based on evolutionary biology and psychology.

Social learning promotes institutions for governing the commons.

Theoretical and empirical research highlights the role of punishment in promoting collaborative efforts. However, both the emergence and the stability of costly punishment are problematic issues. It is not clear how punishers can invade a society of defectors by social learning or natural selection, or how second-order free-riders (who contribute to the joint effort but not to the sanctions) can be prevented from drifting into a coercion-based regime and subverting cooperation. Here we compare the prevailing model of peer-punishment with pool-punishment, which consists in committing resources, before the collaborative effort, to prepare sanctions against free-riders. Pool-punishment facilitates the sanctioning of second-order free-riders, because these are exposed even if everyone contributes to the common good. In the absence of such second-order punishment, peer-punishers do better than pool-punishers; but with second-order punishment, the situation is reversed. Efficiency is traded for stability. Neither other-regarding tendencies or preferences for reciprocity and equity, nor group selection or prescriptions from higher authorities, are necessary for the emergence and stability of rudimentary forms of sanctioning institutions regulating common pool resources and enforcing collaborative efforts.

Evolution of extortion in Iterated Prisoner's Dilemma games.

Iterated games are a fundamental component of economic and evolutionary game theory. They describe situations where two players interact repeatedly and have the ability to use conditional strategies that depend on the outcome of previous interactions, thus allowing for reciprocation. Recently, a new class of strategies has been proposed, so-called "zero-determinant" strategies. These strategies enforce a fixed linear relationship between one's own payoff and that of the other player. A subset of those strategies allows "extortioners" to ensure that any increase in one player's own payoff exceeds that of the other player by a fixed percentage. Here, we analyze the evolutionary performance of this new class of strategies. We show that in reasonably large populations, they can act as catalysts for the evolution of cooperation, similar to tit-for-tat, but that they are not the stable outcome of natural selection. In very small populations, however, extortioners hold their ground. Extortion strategies do particularly well in coevolutionary arms races between two distinct populations. Significantly, they benefit the population that evolves at the slower rate, an example of the so-called "Red King" effect. This may affect the evolution of interactions between host species and their endosymbionts.

Games of corruption: how to suppress illegal logging.

Corruption is one of the most serious obstacles for ecosystem management and biodiversity conservation. In particular, more than half of the loss of forested area in many tropical countries is due to illegal logging, with corruption implicated in a lack of enforcement. Here we study an evolutionary game model to analyze the illegal harvesting of forest trees, coupled with the corruption of rule enforcers. We consider several types of harvesters, who may or may not be committed towards supporting an enforcer service, and who may cooperate (log legally) or defect (log illegally). We also consider two types of rule enforcers, honest and corrupt: while honest enforcers fulfill their function, corrupt enforcers accept bribes from defecting harvesters and refrain from fining them. We report three key findings. First, in the absence of strategy exploration, the harvester-enforcer dynamics are bistable: one continuum of equilibria consists of defecting harvesters and a low fraction of honest enforcers, while another consists of cooperating harvesters and a high fraction of honest enforcers. Both continua attract nearby strategy mixtures. Second, even a small rate of strategy exploration removes this bistability, rendering one of the outcomes globally stable. It is the relative rate of exploration among enforcers that then determines whether most harvesters cooperate or defect and most enforcers are honest or corrupt, respectively. This suggests that the education of enforcers, causing their more frequent trialing of honest conduct, can be a potent means of curbing corruption. Third, if information on corrupt enforcers is available, and players react opportunistically to it, the domain of attraction of cooperative outcomes widens considerably. We conclude by discussing policy implications of our results.

The take-it-or-leave-it option allows small penalties to overcome social dilemmas.

Self-interest frequently causes individuals engaged in joint enterprises to choose actions that are counterproductive. Free-riders can invade a society of cooperators, causing a tragedy of the commons. Such social dilemmas can be overcome by positive or negative incentives. Even though an incentive-providing institution may protect a cooperative society from invasion by free-riders, it cannot always convert a society of free-riders to cooperation. In the latter case, both norms, cooperation and defection, are stable: To avoid a collapse to full defection, cooperators must be sufficiently numerous initially. A society of free-riders is then caught in a social trap, and the institution is unable to provide an escape, except at a high, possibly prohibitive cost. Here, we analyze the interplay of (a) incentives provided by institutions and (b) the effects of voluntary participation. We show that this combination fundamentally improves the efficiency of incentives. In particular, optional participation allows institutions punishing free-riders to overcome the social dilemma at a much lower cost, and to promote a globally stable regime of cooperation. This removes the social trap and implies that whenever a society of cooperators cannot be invaded by free-riders, it will necessarily become established in the long run, through social learning, irrespective of the initial number of cooperators. We also demonstrate that punishing provides a "lighter touch" than rewarding, guaranteeing full cooperation at considerably lower cost.

Moral assessment in indirect reciprocity.

Indirect reciprocity is one of the mechanisms for cooperation, and seems to be of particular interest for the evolution of human societies. A large part is based on assessing reputations and acting accordingly. This paper gives a brief overview of different assessment rules for indirect reciprocity, and studies them by using evolutionary game dynamics. Even the simplest binary assessment rules lead to complex outcomes and require considerable cognitive abilities.

Exploration dynamics in evolutionary games.

Evolutionary game theory describes systems where individual success is based on the interaction with others. We consider a system in which players unconditionally imitate more successful strategies but sometimes also explore the available strategies at random. Most research has focused on how strategies spread via genetic reproduction or cultural imitation, but random exploration of the available set of strategies has received less attention so far. In genetic settings, the latter corresponds to mutations in the DNA, whereas in cultural evolution, it describes individuals experimenting with new behaviors. Genetic mutations typically occur with very small probabilities, but random exploration of available strategies in behavioral experiments is common. We term this phenomenon "exploration dynamics" to contrast it with the traditional focus on imitation. As an illustrative example of the emerging evolutionary dynamics, we consider a public goods game with cooperators and defectors and add punishers and the option to abstain from the enterprise in further scenarios. For small mutation rates, cooperation (and punishment) is possible only if interactions are voluntary, whereas moderate mutation rates can lead to high levels of cooperation even in compulsory public goods games. This phenomenon is investigated through numerical simulations and analytical approximations.

Evolution of indirect reciprocity.

Natural selection is conventionally assumed to favour the strong and selfish who maximize their own resources at the expense of others. But many biological systems, and especially human societies, are organized around altruistic, cooperative interactions. How can natural selection promote unselfish behaviour? Various mechanisms have been proposed, and a rich analysis of indirect reciprocity has recently emerged: I help you and somebody else helps me. The evolution of cooperation by indirect reciprocity leads to reputation building, morality judgement and complex social interactions with ever-increasing cognitive demands.

Incentives and opportunism: from the carrot to the stick.

Cooperation in public good games is greatly promoted by positive and negative incentives. In this paper, we use evolutionary game dynamics to study the evolution of opportunism (the readiness to be swayed by incentives) and the evolution of trust (the propensity to cooperate in the absence of information on the co-players). If both positive and negative incentives are available, evolution leads to a population where defectors are punished and players cooperate, except when they can get away with defection. Rewarding behaviour does not become fixed, but can play an essential role in catalysing the emergence of cooperation, especially if the information level is low.

The competition of assessment rules for indirect reciprocity.

Indirect reciprocity is one of the basic mechanisms to sustain mutual cooperation. Beneficial acts are returned, not by the recipient, but by third parties. Indirect reciprocity is based on reputation and status: it pays to provide help because this makes one more likely to receive help in turn. The mechanism depends on knowing the past behavior of other players, and assessing that behavior. There are many different systems of assessing other individuals, which can be interpreted as rudimentary moral systems (i.e. views on what is 'good' or 'bad'). In this paper, we describe the competition of some of the leading assessment rules called SUGDEN and KANDORI by analytic methods. We show that the sterner rule KANDORI has a slight advantage in the sense that KANDORI-players have more chance to earn higher payoff than SUGDEN-players in the presence of unconditional altruists. On the other hand, we see that the unconditional altruists are eliminated in the long run and that stable polymorphisms of KANDORI and SUGDEN can subsist, but that a moral consensus is realized even in those polymorphic states: all players' images are the same in each observer's eyes.

Partners or rivals? Strategies for the iterated prisoner's dilemma.

Within the class of memory-one strategies for the iterated Prisoner's Dilemma, we characterize partner strategies, competitive strategies and zero-determinant strategies. If a player uses a partner strategy, both players can fairly share the social optimum; but a co-player preferring an unfair solution will be penalized by obtaining a reduced payoff. A player using a competitive strategy never obtains less than the co-player. A player using a zero-determinant strategy unilaterally enforces a linear relation between the two players' payoffs. These properties hold for every strategy used by the co-player, whether memory-one or not.

Punishment and reputation in spatial public goods games.

The puzzle of the emergence of cooperation between unrelated individuals is shared across diverse fields of behavioural sciences and economics. In this article we combine the public goods game originating in economics with evolutionary approaches traditionally used in biology. Instead of pairwise encounters, we consider the more complex case of groups of three interacting individuals. We show that territoriality is capable of promoting cooperative behaviour, as in the case of the Prisoner's Dilemma. Moreover, by adding punishment opportunities, the readiness to cooperate is greatly enhanced and asocial strategies can be largely suppressed. Finally, as soon as players carry a reputation for being willing or unwilling to punish, highly cooperative and fair outcomes are achieved. This group-beneficial result is obtained, intriguingly, by making individuals more likely to exploit their co-players if they can get away with it. Thus, less-cooperative individuals make more-cooperative societies.