“…Studies of alpha male descent in natural populations of primates have found that it occurs for many possible reasons, including the alpha male being no longer physically capable of staving off challenger males, alpha males losing coalitionary support, the immigration of more dominant individuals into the social group, or the sexual maturation of younger, more dominant individuals (O'Shea, 1976;Perry, 1998;Uehara et al, 1994). It is highly metabolically costly for alpha males to consistently defend their dominance status and territory through physical fighting (Briffa & Sneddon, 2007;Castro, Ros, Becker, & Oliveira, 2006;Rohwer & Ewald, 1981) and other behaviours such as scent marking (Gosling, Roberts, Thornton, & Andrew, 2000). Dominant alpha males of many species also have higher levels of testosterone and cortisol that may be physiologically damaging (Gesquiere et al, 2011;Higham, Heistermann, & Maestripieri, 2012;Mendonça-Furtado et al, 2014;Sapolsky, 2005).…”