title: Long-distance communication facilitates cooperation among wild spotted hyaenas,Crocuta crocuta.
abstract: Calls that catalyse group defence, as in the mobbing of predators, appear to facilitate cooperation by recruiting receivers to act collectively. However, even when such signals reliably precede cooperative behaviour, the extent to which the calls function as recruitment signals is unclear. Calls might simply arouse listeners' attention, setting off a cascade of independent responses to the threat. By contrast, they might convey information, for example, about signaller identity and the nature of a threat that affects receivers' decisions to participate. We explored this distinction by investigating a possible long-distance recruitment call used by spotted hyaenas. These social carnivores live in fission-fusion clans and individuals disperse widely within their territories. Putative recruitment calls must therefore attract receivers that are distant from the inciting threat and free to opt out of risky collective aggression. Hyaenas compete with lions over food, and neighbouring clans sometimes engage in violent border clashes. These high-stakes contests are decided based on numerical asymmetries, so hyaenas can only protect critical resources if the dispersed clan can converge quickly at conflict sites. We recorded and analysed whoop bouts produced in multiple contexts and found that bouts produced in response to signs of lion-hyaena conflict had shorter inter-whoop intervals than spontaneous 'display' bouts. In subsequent field playback experiments, resting hyaenas were significantly more likely to move in response to 'recruitment' bouts with shortened intervals than to otherwise identical 'display' bouts. Whereas only stimulus type predicted movement, lower-ranked subjects responded most quickly, perhaps because their feeding opportunities depend on arriving early at any kill site. Results demonstrate that hyaenas possess a signal that can reliably recruit allies across long distances, despite moderating effects of individual circumstances on the strength of receivers' responses.
authors.affiliation: Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, U.S.A.
authors.forename: Andrew S
authors.initials: AS
authors.lastname: Gersick
authors.affiliation: Department of Biology, University of Pennsylvania, Philadelphia, PA, U.S.A.
authors.forename: Dorothy L
authors.initials: DL
authors.lastname: Cheney
authors.affiliation: Philadelphia, PA, U.S.A.
authors.forename: Jennifer M
authors.initials: JM
authors.lastname: Schneider
authors.affiliation: Institute for Research in Cognitive Science, Department of Psychology, University of Pennsylvania, Philadelphia, PA, U.S.A.
authors.forename: Robert M
authors.initials: RM
authors.lastname: Seyfarth
authors.affiliation: Department of Zoology, Michigan State University, East Lansing, MI, U.S.A.
authors.forename: Kay E
authors.initials: KE
authors.lastname: Holekamp
keywords: Crocuta crocuta; alarm signal; animal communication; cooperative defence; recruitment; spotted hyaena