Behavioral Ecology and Sociobiology

, Volume 63, Issue 10, pp 1495–1503 | Cite as

Behavioural trait assortment in a social network: patterns and implications

  • Darren P. CroftEmail author
  • Jens Krause
  • Safi K. Darden
  • Indar W. Ramnarine
  • Jolyon J. Faria
  • Richard James
Original Paper


The social fine structure of a population plays a central role in ecological and evolutionary processes. Whilst many studies have investigated how morphological traits such as size affect social structure of populations, comparatively little is known about the influence of behaviours such as boldness and shyness. Using information on social interactions in a wild population of Trinidadian guppies (Poecilia reticulata), we construct a social network. For each individual in the network, we quantify its behavioural phenotype using two measures of boldness, predator inspection tendency, a repeatable and reliably measured behaviour well studied in the context of co-operation, and shoaling tendency. We observe striking heterogeneity in contact patterns, with strong ties being positively assorted and weak ties negatively assorted by our measured behavioural traits. Moreover, shy fish had more network connections than bold fish and these were on average stronger. In other words, social fine structure is strongly influenced by behavioural trait. We assert that such structure will have implications for the outcome of selection on behavioural traits and we speculate that the observed positive assortment may act as an amplifier of selection contributing to the maintenance of co-operation during predator inspection.


Behavioural phenotype Co-operation Evolutionary graph theory Guppy Personality Poecilia reticulata Predator inspection 



We would like to thank P. Thomas, M. Botham, J. Dyer, J. Ward and C. Piyapong for assistance with data collection and Graeme Ruxton and two anonymous referees for valuable comments on a previous version of this manuscript. Funding was provided to DPC by NERC (NE/E001181/1) and JK by the EPSRC (GR/T11241/01(P)).


  1. Aktipis CA (2004) Know when to walk away: contingent movement and the evolution of cooperation. J Theor Biol 231:249–260PubMedCrossRefGoogle Scholar
  2. Budaev SV (1997) “Personality” in the guppy (Poecilia reticulata): a correlational study of exploratory behavior and social tendency. J Comp Psychol 111:399–411CrossRefGoogle Scholar
  3. Cairns SJ, Schwager SJ (1987) A comparison of association indexes. Anim Behav 35:1454–1469CrossRefGoogle Scholar
  4. Conradt L (1998) Could asynchrony in activity between the sexes cause intersexual social segregation in ruminants? Proc R Soc Lond B Biol Sci 265:1359–1363CrossRefGoogle Scholar
  5. Croft DP, Albanese B, Arrowsmith BJ, Botham M, Webster M, Krause J (2003a) Sex biased movement in the guppy (Poecilia reticulata). Oecologia 137:62–68PubMedCrossRefGoogle Scholar
  6. Croft DP, Arrowsmith BJ, Bielby J, Skinner K, White E, Couzin ID, Magurran AE, Ramnarine I, Krause J (2003b) Mechanisms underlying shoal composition in the Trinidadian guppy (Poecilia reticulata). Oikos 100:429–438CrossRefGoogle Scholar
  7. Croft DP, Krause J, James R (2004) Social networks in the guppy (Poecilia reticulata). Proc R Soc Lond B Biol Sci 271:S516–S519CrossRefGoogle Scholar
  8. Croft DP, James R, Ward AJW, Botham MS, Mawdsley D, Krause J (2005) Assortative interactions and social networks in fish. Oecologia 143:211–219PubMedCrossRefGoogle Scholar
  9. Croft DP, James R, Thomas POR, Hathaway C, Mawdsley D, Laland KN, Krause J (2006a) Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata). Behav Ecol Sociobiol 59:644–650CrossRefGoogle Scholar
  10. Croft DP, Morrell LJ, Wade AS, Piyapong C, Ioannou CC, Dyer JRG, Chapman BB, Yan W, Krause J (2006b) Predation risk as a driving force for sexual segregation: a cross-population comparison. Am Nat 167:867–878CrossRefGoogle Scholar
  11. Croft DP, James R, Krause J (2008) Exploring animal social networks. Princeton University Press, Princeton, NJGoogle Scholar
  12. Darden SK, Croft DP (2008) Male harassment drives females to alter habitat use and leads to segregation of the sexes. Biol Lett 4:449–451PubMedCrossRefGoogle Scholar
  13. Dugatkin LA (1988) Do guppies play tit for tat during predator inspection visits? Behav Ecol Sociobiol 23:395–399CrossRefGoogle Scholar
  14. Dugatkin LA (1992) Tendency to inspect predators predicts mortality risk in the guppy (Poecilia reticulata). Behav Ecol 3:124–127CrossRefGoogle Scholar
  15. Dugatkin LA, Alfieri M (1991) Guppies and the tit-for-tat strategy - preference based on past interaction. Behav Ecol Sociobiol 28:243–246CrossRefGoogle Scholar
  16. Dugatkin LA, Wilson DS (2000) Assortative interactions and the evolution of cooperation during predator inspection in guppies (Poecilia reticulata). Evol Ecol Res 2:761–767Google Scholar
  17. Godin JGJ, Davis SA (1995) Who dares, benefits: predator approach behaviour in the guppy (Poecilia reticualta) deters predator pursuit. Proc R Soc Lond B Biol Sci 259:193–200CrossRefGoogle Scholar
  18. Gosling SD (2001) From mice to men: what can we learn about personality from animal research? Psychol Bull 127:45–86PubMedCrossRefGoogle Scholar
  19. Huntingford F (1976) The relationship between anti-predator behaviour and aggression among conspecifics in the three-spined stickleback, Gasterosteus aculeatus. Anim Behav 24:245–260CrossRefGoogle Scholar
  20. James R, Croft DP, Krause J (2009) Potential banana skins in animal social network analysis. Behav Ecol Sociobiol 63:989–997CrossRefGoogle Scholar
  21. Krause J, Ruxton GD (2002) Living in groups. Oxford University Press, OxfordGoogle Scholar
  22. Krause J, Butlin RK, Peuhkuri N, Pritchard VL (2000) The social organization of fish shoals: a test of the predictive power of laboratory experiments for the field. Biol Rev 75:477–501PubMedGoogle Scholar
  23. Krause J, Croft DP, James R (2007) Social network theory in the behavioural sciences: potential applications. Behav Ecol Sociobiol 62:15–27CrossRefGoogle Scholar
  24. Krause J, Lusseau D, James R (2009) Animal social networks: an introduction. Behav Ecol Sociobiol 63:967–973CrossRefGoogle Scholar
  25. Külling D, Milinski M (1992) Size-dependent predation risk and partner quality in predator inspection of sticklebacks. Anim Behav 44:949–955CrossRefGoogle Scholar
  26. Landeau L, Terborgh J (1986) Oddity and the confusion effect in predation. Anim Behav 34:1372–1380CrossRefGoogle Scholar
  27. Magurran AE (1993) Individual differences and alternative behaviours. In: Pitcher TJ (ed) Behaviour of teleost fishes, 2nd edn. Chapman & Hall, LondonGoogle Scholar
  28. Magurran AE, Higham A (1988) Information transfer across fish shoals under predator threat. Ethology 78:153–158CrossRefGoogle Scholar
  29. Maynard Smith J (1982) Evolution and the theory of games. Cambridge University Press, CambridgeGoogle Scholar
  30. Milinski M (1987) Tit-for-Tat in sticklebacks and the evolution of cooperation. Nature 325:433–435PubMedCrossRefGoogle Scholar
  31. Milinski M, Luthi JH, Eggler R, Parker GA (1997) Cooperation under predation risk: experiments on costs and benefits. Proc R Soc Lond B Biol Sci 264:831–837CrossRefGoogle Scholar
  32. Milinski M, Pfluger D, Külling D, Kettler R (1990) Do sticklebacks cooperate repeatedly in reciprocal pairs? Behav Ecol Sociobiol 27:17–21CrossRefGoogle Scholar
  33. Newman MEJ (2003) Mixing patterns in networks. Phys Rev E 67:art. no.-026126Google Scholar
  34. Ohguchi O (1978) Experiments on the selection against colour oddity of water fleas by three-spined stickelbacks. Z Tierpsychol 47:254–267Google Scholar
  35. Ohtsuki H, Hauert C, Lieberman E, Nowak MA (2006) A simple rule for the evolution of cooperation on graphs and social networks. Nature 441:502–505PubMedCrossRefGoogle Scholar
  36. Pike TW, Samanta M, Lindström J, Royle NJ (2008) Behavioural phenotype affects interactions in a social network. Proc R Soc Lond B Biol Sci 275:2515–2520CrossRefGoogle Scholar
  37. Pitcher TJ, Green DA, Magurran AE (1986) Dicing with death—predator inspection behaviour in minnow shoals. J Fish Biol 28:439–448CrossRefGoogle Scholar
  38. Reale D, Reader SM, Sol D, McDougall PT, Dingemanse NJ (2007) Integrating animal temperament within ecology and evolution. Biol Rev 82:291–318PubMedCrossRefGoogle Scholar
  39. Roberts SGB, Wilson R, Fedurek P, Dunbar RIM (2008) Individual differences and personal social network size and structure. Personality Individ Differ 44:954–964CrossRefGoogle Scholar
  40. Ruckstuhl KE (1999) To synchronise or not to synchronise: a dilemma for young bighorn males? Behaviour 136:805–818CrossRefGoogle Scholar
  41. Santos FC, Pacheco JM, Lenaerts T (2006) Cooperation prevails when individuals adjust their social ties. PLoS Comput Biol 2:1284–1291CrossRefGoogle Scholar
  42. Sih A, Bell A, Johnson JC (2004) Behavioral syndromes: an ecological and evolutionary overview. Trends Ecol Evol 19:372–378PubMedCrossRefGoogle Scholar
  43. Szulkin M, Dawidowicz P, Dodson SI (2006) Behavioural uniformity as a response to cues of predation risk. Anim Behav 71:1013–1019CrossRefGoogle Scholar
  44. Theodorakis CW (1989) Size segregation and the effects of oddity on predation risk in minnow schools. Anim Behav 38:496–502CrossRefGoogle Scholar
  45. Thomas POR, Croft DP, Morrell LJ, Davis A, Faria JJ, Dyer JRG, Piyapong C, Ramnarine I, Ruxton GD, Krause J (2008) Does defection during predator inspection affect social structure in wild shoals of guppies? Anim Behav 75:43–53CrossRefGoogle Scholar
  46. Wey T, Blumstein DT, Shen W, Jordan F (2008) Social network analysis of animal behaviour: a promising tool for the study of sociality. Anim Behav 75:333–344CrossRefGoogle Scholar
  47. Wilson EO (1975) Sociobiology: the new synthesis. Harvard University Press, Cambridge, MAGoogle Scholar
  48. Wilson DS (1998) Adaptive individual differences within single populations. Philos Trans R Soc B-Biol Sci 353:199–205CrossRefGoogle Scholar
  49. Wilson DS, Dugatkin LA (1997) Group selection and assortative interactions. Am Nat 149:336–351CrossRefGoogle Scholar
  50. Wilson DS, Coleman K, Clark AB, Biederman L (1993) Shy bold continuum in pumpkinseed sunfish (Lepomis gibbosus): an ecological study of a psychological trait. J Comp Psychol 107:250–260CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2009

Authors and Affiliations

  • Darren P. Croft
    • 1
    • 2
    Email author
  • Jens Krause
    • 3
  • Safi K. Darden
    • 1
    • 2
  • Indar W. Ramnarine
    • 4
  • Jolyon J. Faria
    • 3
  • Richard James
    • 5
  1. 1.School of Biological Sciences, College of Natural SciencesBangor UniversityBangorUK
  2. 2.Centre for Research in Animal Behaviour, School of PsychologyUniversity of ExeterExeterUK
  3. 3.Institute of Integrative and Comparative BiologyUniversity of LeedsLeedsUK
  4. 4.Department of Life SciencesThe University of the West Indies, St. AugustineSt. AugustineTrinidad & Tobago
  5. 5.Department of PhysicsUniversity of BathBathUK

Personalised recommendations