“…For example, they can identify individual conspecifics, and distinguish kin from nonkin, on the basis of their long-distance ‘whoop’ vocalizations, and whoops also convey information about the caller’s age, sex and motivational state (Benson-Amram, Heinen, Dryer, & Holekamp, 2011; East & Hofer, 1991a, 1991b; Gersick et al, 2015; Holekamp et al, 1999; Theis, Greene, Benson-Amram, & Holekamp, 2007). Hyaenas also have a keen olfactory sense; each clan has a unique scent signature, mediated in part by volatile products of metabolism in the symbiotic microbes inhabiting the hyaenas’ scent glands (Hofer, East, Sammang, & Dehnhard, 2001; Theis, Schmidt, & Holekamp, 2012, Theis et al, 2013), and hyaenas can distinguish scents of their clan-mates from those of hyaenas from other clans (Theis, 2007).…”