Abstract

When the benefits of interacting with out-group members exceed the associated costs, social groups may be expected to be tolerant towards each other. However, in many species exhibiting intergroup tolerance, the nature of benefits gained from intergroup encounters remains unclear. We investigated the potential costs and benefits associated with intergroup associations in bonobos, a species with varying degrees of intergroup tolerance, by testing whether these associations conferred energetic benefits to participants under different socioecological contexts and whether the consequences of these associations substantially differed from within-group competition. We used measures of socioecological factors (fruit abundance and group size), feeding and ranging behaviors, and a physiological marker of energy balance (urinary c-peptide of insulin) collected over a 19-month period from two neighboring wild communities in the Kokolopori Bonobo Reserve, Democratic Republic of the Congo. We found that intergroup associations were not related to individuals’ energy balance, but they were related to variations in individuals’ ranging and feeding behavior. Specifically, bonobos traveled longer distances, visited larger fruit patches, and increased the time spent feeding on fruits on days they associated with the neighboring group. These adaptations in feeding behavior may be strategies to offset the energetic costs of increased travel distances. In the absence of obvious energetic benefits and with clear strategies employed to offset energetic costs, it is likely that intergroup associations in bonobos provide benefits unrelated to energy acquisition, such as social benefits. Our study sheds light on the potential incentives promoting social networks to extend beyond and across groups in a tolerant species.Significance statementIntergroup encounters can be energetically costly due to increased competition over resources. Yet, some species associate with out-group individuals for extended periods of time when the benefits of participating in these associations exceed the potential costs. Bonobos, a species exhibiting intergroup tolerance, modified their feeding behavior during intergroup associations by feeding on larger fruit patches and increasing their time spent feeding on fruits, likely to offset energetic costs of increased travel distances. As results, individuals’ energy balance was not related with intergroup associations. The employment of such strategies in addition to the absence of clear energetic benefits suggests that intergroup associations in bonobos provide social rather than ecological benefits.

Highlights

  • Reactions to out-group members in group-living species range from agonistic to tolerant and even affiliativeBehav Ecol Sociobiol (2021) 75: 2(polydomous ants, Iridomyrmex purpureus: Robinson and Barker 2017; humans and great apes: Pisor and Surbeck 2019), with some species such as humans spanning the full range of potential reactions (Kelly 2005; Fry 2012)

  • We investigated whether intercommunity associations in bonobos conferred clear energetic benefits to individuals, as measured by variation in urinary c-peptide of insulin (UCP) levels

  • We found indications that associations carried energetic costs due to longer travel distances and that the bonobos actively offset these costs by visiting larger fruit patches and increasing the time spent feeding on fruits on association days

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Summary

Introduction

(polydomous ants, Iridomyrmex purpureus: Robinson and Barker 2017; humans and great apes: Pisor and Surbeck 2019), with some species such as humans spanning the full range of potential reactions (Kelly 2005; Fry 2012) The nature of these interactions is influenced by a variety of socioecological factors including the availability, distribution, and location of feeding resources (spotted hyenas, C. crocuta: Boydston et al 2001; coal tits, Parus ater: Brotons and Herrando 2003; non-human primates review: Koenig 2002; Brown 2013; banded mongooses, M. mungo: Furrer et al 2011; Verreaux’s sifakas, Propithecus verreauxi: Koch et al 2016; polydomous ants, I. purpureus; and humans: Robinson and Barker 2017) and mating opportunities (humpback whales, Megaptera novaeangliae: Clapham et al 1992; chacma baboons, Papio cynocephalus ursinus: Kitchen et al 2004), as well as the relative sizes of the interacting groups (wood ants, Formica xerophila: Tanner 2006; lions, Panthera leo: Mosser and Packer 2009). Such costs generally originate from higher competition over food resources due to a larger number of co-feeding individuals (Chapman and Valenta 2015; Markham et al 2015), which may result in increased travel distances to find sufficient food (chimpanzees, Pan troglodytes and spider monkeys, Ateles geoffroyi: Chapman et al 1995; red colobus monkeys, Procolobus badius: Gillespie and Chapman 2001), increased time individuals allocate to feeding (yellow baboons, P. cynocephalus: Muruthi et al 1991; black-and-white colobus, Colobus polykomos: Dasilva 1992), and higher number and/or size of feeding patches visited (howler monkeys, Alouatta palliata: Leighton and Leighton 1982; white-throated sparrows, Zonotrichia albicollis: Pearson 1989; muriquis, Brachyteles arachnoides: Strier 1989)

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