“…Multiple mechanisms have been proposed to achieve assortment (see Nowak, 2006, for a review), such as: (i) the existence of an underlying spatial structure combined with limited dispersal, also known as population viscosity, which clusters together parents and offspring (Hamilton, 1964;Lieberman et al, 2005;Nowak and May, 1992;Nowak et al, 2010a;Pfeiffer and Bonhoeffer, 2003;Tarnita et al, 2009b;West et al, 2006); (ii) structure arising from the properties of the individuals such as adhesion, speed of movement or perception of environmental cues (Avilés, 2002;Garcia et al, 2014Garcia et al, , 2015; (iii) recognition of like or discrimination of non-like based on various cues such as kinship, phenotypic similarity, signaling or shared preferences (Antal et al, 2009;Mehdiabadi et al, 2006;Ostrowski et al, 2008;Pacheco et al, 2006;Rousset and Roze, 2007;Stefanic et al, 2015;Tarnita et al, 2009a;Traulsen and Nowak, 2007;Vos and Velicer, 2009); (iv) reciprocity, which relies on either memory or reputation to ensure that future interactions are informed by previous ones (Axelrod and Hamilton, 1981;Nowak andSigmund, 1989, 2005;Trivers, 1971); or (v) multilevel selection Wilson, 1975), which considers selective pressures both at the individual Glossary Cooperation Costly behavior that confers fitness benefits on same-species recipients. Defection (free-riding) Absence of the cooperative behavior coupled with exploitation of its benefits.…”