Abstract

Predation pressure has long been considered a leading explanation of colonies, where close neighbors may reduce predation via dilution, alarming or group predator attacks. Attacking predators may be costly in terms of energy and survival, leading to the question of how neighbors contribute to predator deterrence in relationship to each other. Two hypotheses explaining the relative efforts made by neighbors are byproduct-mutualism, which occurs when breeders inadvertently attack predators by defending their nests, and reciprocity, which occurs when breeders deliberately exchange predator defense efforts with neighbors. Most studies investigating group nest defense have been performed with birds. However, colonial fish may constitute a more practical model system for an experimental approach because of the greater ability of researchers to manipulate their environment. We investigated in the colonial fish, Neolamprologus caudopunctatus, whether prospecting pairs preferred to breed near conspecifics or solitarily, and how breeders invested in anti-predator defense in relation to neighbors. In a simple choice test, prospecting pairs selected breeding sites close to neighbors versus a solitary site. Predators were then sequentially presented to the newly established test pairs, the previously established stimulus pairs or in between the two pairs. Test pairs attacked the predator eight times more frequently when they were presented on their non-neighbor side compared to between the two breeding sites, where stimulus pairs maintained high attack rates. Thus, by joining an established pair, test pairs were able to reduce their anti-predator efforts near neighbors, at no apparent cost to the stimulus pairs. These findings are unlikely to be explained by reciprocity or byproduct-mutualism. Our results instead suggest a commensal relationship in which new pairs exploit the high anti-predator efforts of established pairs, which invest similarly with or without neighbors. Further studies are needed to determine the scope of commensalism as an anti-predator strategy in colonial animals.

Highlights

  • Predation of offspring has long been stressed as one of the major factors affecting the fitness of breeders [1,2]

  • For the analyses of the predator presentation, we focused on attacks because they were the most unambiguous behavior identified as nest defense and intruder deterrence

  • When we presented predators to test pairs that had settled near the stimulus pair, their attack rates were approximately eight times higher at the far side of the breeding site than at the side neighboring the stimulus pair (Fig. 2; position A vs. B: N = 11, Wilcoxon Z = –2.93, p = 0.003)

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Summary

Introduction

Predation of offspring has long been stressed as one of the major factors affecting the fitness of breeders [1,2] This factor has spawned many studies about behavioral strategies of nest defense, which often comprise direct attacks or approaches by breeders to drive away predators. In tree swallows Tachycineta tricolor for example, multiple neighbors mobbed predators placed near their nests, thereby increasing the intensity of predator defense per nest [15]. This effect was shown in colonial Montagu’s harriers Circus pygargus in which the probability of the predator model being attacked increased with group size [16]. The individual rate of high-risk diving-attacks decreased with group size, suggesting that breeding near close neighbors reduces the individual net costs of defense [16]

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