Abstract

Territorial animals often use signals to advertise territorial occupancy within their larger home ranges. Songbirds are among the best-studied territorial signaling taxa, and when competitors start singing during a territorial intrusion, residents usually show elevated spatial and vocal responses. These responses could be used by intruders and distant eavesdroppers to predict future responses or to compare responses across competitors. Yet, the extent to which responses of a resident to a territorial intrusion predict its future responses and its overall spatial behavior (home range) within a neighborhood is less well understood. We used wild great tits (Parus major) as a model species in repeated song playback trials, simulating territorial intrusions combined with radio-tracking before and during playback trials. The time spent close to the loudspeaker in response to an initial simulated intrusion predicted the same response variable during a second simulated intrusion on the next day, whereas singing activity during the first simulated intrusion did not predict singing during the second simulated intrusion. We also show that more explorative males (as determined by a novel environment test) and males with smaller home ranges sang more and spent more time near the loudspeaker in response to both simulated intrusions. Thus, by probing residents, intruders can obtain reliable information about subsequent response probabilities, while eavesdroppers from a distance, who can use auditory information only, would not receive sufficient predictive information. Our findings also suggest that males with larger home ranges are more tolerant toward intruders, which could reflect a trade-off between tendencies to respond strongly and to range widely. The lack of predictability of singing activity with regard to responses to future intrusions might explain why territorial animals continuously exchange vocal signals and regularly foray into neighboring territories, as a way to obtain regular information updates.Significance StatementAnimals use experience from interactions with conspecifics in their future decision making, such as mate choice and strategies for conflict resolution. The value of such information depends in part on the predictability of the future behavior of that conspecific. In songbirds, territorial individuals respond to intruders by approach and signaling. Here, we tested in radio-tagged great tits (Parus major) if territorial responses are predictable and are affected by individual and environmental factors. We show that the time spent near the simulated intruder was more predictable than singing activity and that birds with larger home ranges showed weaker responses. These findings suggest that information based on such spatial responses is more useful for future decision making, as compared to vocal information, and that distant eavesdroppers will thus receive less reliable information. Limited predictability may explain why territorial animals continuously exchange vocal signals and foray into neighboring territories, providing opportunities for regular information updates.

Highlights

  • Many animals use signals to defend resources, such as mates, feeding sites, or territories

  • We show that the time spent near the simulated intruder was more predictable than singing activity and that birds with larger home ranges showed weaker responses

  • Using a combination of radio-tracking and playback trials that simulated territorial intrusions, we show that territory owners’ time spent close to a simulated intruder was predicted by this same measure during a prior intrusion

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Summary

Introduction

Many animals use signals to defend resources, such as mates, feeding sites, or territories. Competitors frequently trespass into others’ territories without necessarily challenging the resident, presumably to explore and assess the territory or resident (Naguib et al 2008; Snijders and Naguib 2017) Such intrusions may be part of a process in which space gets partitioned, for example, by testing rivals’ alertness and responsiveness (Stamps and Krishnan 2001), potentially leading to continuous small changes in initial territory sizes and boundary locations. Such intrusions are known to occur after territories appear to be fully established (Hanski and Haila 1988; Chandler et al 1994, 1997; Pitcher and Stutchbury 2000; Naguib et al 2001; Bircher et al 2020), likely to continually update information about neighboring conspecifics

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