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Use of social network models to understand play partner choice strategies in three primate species

  • Meredith C. LutzEmail author
  • Jonah Ratsimbazafy
  • Peter G. Judge
Special Feature: Original Article Social networks analysis in primates, a multilevel perspective


Although play is seen in many species, its evolutionary function is still largely unknown. Several relevant, proposed hypotheses (such as the training for the unexpected, self-assessment, social skills, and dominance hierarchy hypotheses) make predictions about how animals should optimally choose their play partners based on their familiarity or other demographic variables. We used a social network approach to analyze focal sample data on brown capuchins (Cebus apella), hamadryas baboons (Papio hamadryas), and diademed sifaka (Propithecus diadema) to understand how these species choose their play partners with respect to demographic variables. Using exponential random graph models (ERGMs), we found that sifaka and capuchins generally tended to play with animals who were similar to them. The baboons were only sensitive to age differences in the formation of strong play relationships. Our data most strongly support the training for the unexpected hypothesis, as according to predictions all species preferred to play with animals who were their close social partners, decreasing the possibility of cheating during play. Through the first application (to our knowledge) of ERGMs to primate behavior, we were able to compare the effects of many demographic variables on the complex, interdependent social structure of primates. Applying this tool to additional groups and species will provide further insight into evolutionary mechanisms of play behavior across taxa.


Social play behavior Training for the unexpected hypothesis Social skills hypothesis Self-assessment hypothesis Exponential random graph models 



We would like to thank Mary Gavitt, Gretchen Long, Amber Hackenberg, and the rest of Bucknell’s animal caretakers for their excellent care of all the monkeys at the Bucknell Animal Behavior Laboratory. We would also like to thank Ravi Goyal, Mathematica, for his advice on the analysis. We would like to thank the entire GERP team in Antananarivo, Madagascar for their help in organizing research permits, and the Maromizaha team in Anevoka for arranging logistics while on site. Data collection in the field was supported by MAMINIRIMA Herinjetovo and RANDIANATOANDRO Heritiana. Additional dominance data was collected by Corinne Leard, Allie Schrock, Melissa Painter, Danielle Antonellis, and Amelia Lautenberg. We also thank Dr. Matthew Silk and two anonymous reviewers for their helpful comments on earlier drafts of the manuscript. Thanks also to Mark Grote for helpful guidance on aspects of the analysis during the review process and to Damien Caillaud, Alexandra McInturf, and Neetha Iyer for helpful discussions on the manuscript revisions. MCL was supported by a Bucknell Program for Undergraduate Research Grant, two Bucknell University Animal Behavior summer research grants, a Bucknell University Statistics summer research grant, and a Bucknell University Presidential Fellowship. The primate facility was supported by Bucknell University.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed.

Supplementary material

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Copyright information

© Japan Monkey Centre and Springer Japan KK, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Animal Behavior ProgramBucknell UniversityLewisburgUSA
  2. 2.Department of MathematicsBucknell UniversityLewisburgUSA
  3. 3.Animal Behavior Graduate GroupUniversity of California, DavisDavisUSA
  4. 4.Le Groupe d’Étude et de Recherche sur les Primates de Madagascar (GERP)AntananarivoMadagascar
  5. 5.Department of Paleontology and AnthropologyUniversity of AntananarivoAntananarivoMadagascar
  6. 6.Department of PsychologyBucknell UniversityLewisburgUSA

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