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Trust and trustworthiness form the basis for continued social and economic interactions, and they are also fundamental for cooperation, fairness, honesty, and indeed for many other forms of prosocial and moral behaviour. However, trust entails risks, and building a trustworthy reputation requires effort. So how did trust and trustworthiness evolve, and under which conditions do they thrive? To find answers, we operationalize trust and trustworthiness using the trust game with the trustor's investment and the trustee's return of the invest