To Bees or Not to Bees: Greater Honeyguides Sometimes Guide Humans to Animals Other Than Bees, but Likely Not as Punishment.

Greater honeyguides () are well known to guide human honey hunters to wild bees' nests in exchange for beeswax as food. Centuries of African Indigenous accounts have intriguingly reported that honeyguides occasionally guide humans to animals other than bees, typically large animals dangerous to humans. This is interpreted by some human cultures as punishment for prior failure to reward the bird, and by others as an altruistic warning behavior. Here, we present quantitative evidence from hundreds of honeyguide-human interactions in Mozambique of greater honeyguides guiding humans to snakes ( = 3) and a dead mammal ( = 1). We show that guiding behavior to these vertebrates was (i) spatially and acoustically analogous to honeyguide behavior when guiding to bees, (ii) did not occur more frequently after not being rewarded with beeswax by humans, and (iii) was rare (3.7% of human-honeyguide interactions in 1 year; 0% in others). We review historical accounts and cultural explanations for this behavior and use these to inform five hypotheses for why honeyguides guide people to nonbee animals. Our field data were most consistent with the hypothesis that guiding to nonbee animals results from a cognitive recall error of spatial information. We suggest that this behavior is unlikely to function as punishment, yet may coincidentally benefit honeyguides over longer timescales by initiating a human cultural interpretation that reinforces human cultural traditions of rewarding honeyguides with beeswax.