Behavioral Ecology and Sociobiology

, Volume 62, Issue 2, pp 289–298 | Cite as

Emergence of increased division of labor as a function of group size

  • Raphaël Jeanson
  • Jennifer H. Fewell
  • Root Gorelick
  • Susan M. Bertram
Original Paper


Empirical evidence suggests that division of labor in insect societies is positively related to group size both within and across taxa. Response threshold models (RTM) have been commonly used to analyze patterns of division of labor. However, these models have been explored empirically and theoretically for only a limited number of tasks, and few studies have examined predictions of the model as colony size and work availability change. We theoretically examine how group size influences division of labor using a fixed response-threshold model. We simultaneously explore how expected by-products of increased colony size, including demand (total work need relative to total work force available) and task number, affect this relationship. Our results indicate that both low demand and high task number positively influence division of labor. We suggest that these changes parallel what is observed within social groups as their size increases, and that, in part, the commonly observed increased division of labor with increasing group size is emergent.


Colony size Division of labor Tasks number Threshold model 



We wish to thank B. Hölldobler and P. Kukuk for their helpful discussions and insights. We thank four anonymous referees for helpful comments regarding the manuscript. This research was supported by National Science Foundation grant number 0446415 awarded to JHF and SMB. RJ was supported by a post-doctoral grant from the Fyssen Foundation.


  1. Anderson C (2001) The adaptive value of inactive foragers and the scout-recruit system in honey bee (Apis mellifera) colonies. Behav Ecol 12:111–119Google Scholar
  2. Anderson C, Ratnieks FLW (1999) Task partitioning in insect societies. I. Effect of colony size on queueing delay and colony ergonomic efficiency. Am Nat 154:521–535PubMedCrossRefGoogle Scholar
  3. Anderson C, McShea D (2001) Individual versus social complexity, with particular reference to ant colonies. Biol Rev 76:211–237PubMedCrossRefGoogle Scholar
  4. Arathi HS, Spivak M (2001) Influence of colony genotypic composition on the performance of hygienic behaviour in the honeybee, Apis mellifera L. Anim Behav 62:57–66CrossRefGoogle Scholar
  5. Beekman M, Sumpter DJT, Ratnieks FLW (2001) Phase transition between disordered and ordered foraging in Pharaoh’s ants. Proc Natl Acad Sci U S A 98:9703–9706PubMedCrossRefGoogle Scholar
  6. Bell G, Mooers AO (1997) Size and complexity among multicellular organisms. Biol J Linn Soc 60:345–363CrossRefGoogle Scholar
  7. Beshers SN, Fewell JH (2001) Models of division of labor in social insects. Ann Rev Entomol 46:413–440CrossRefGoogle Scholar
  8. Bonabeau E, Theraulaz G, Deneubourg J-L (1996) Quantitative study of the fixed threshold model for the regulation of division of labour in insect societies. Proc R Soc Lond B 263:1565–1569CrossRefGoogle Scholar
  9. Bonabeau E, Theraulaz G, Deneubourg JL (1998) Fixed response thresholds and the regulation of division of labor in insect societies. Bull Math Biol 60:753–807CrossRefGoogle Scholar
  10. Bonner JT (1993) Dividing the labor in cells and societies. Curr Sci 64:459–466Google Scholar
  11. Bonner JT (1998) The origins of multicellularity. Integr Biol 1:27–36CrossRefGoogle Scholar
  12. Bonner JT (2004) Perspective: The size-complexity rule. Evolution 58:1883–1890PubMedGoogle Scholar
  13. Boomsma JJ, Ratnieks FLW (1996) Paternity in eusocial hymenoptera. Philos Trans R Soc Lond B 351:947–975CrossRefGoogle Scholar
  14. Bourke AFG (1999) Colony size, social complexity and reproductive conflict in social insects. J Evol Biol 12:245–257CrossRefGoogle Scholar
  15. Cole BJ (1983) Multiple mating and the evolution of social behavior in the Hymenoptera. Behav Ecol Sociobiol 12:191–201CrossRefGoogle Scholar
  16. Darchen R (1964) Biologie des Vespa orientalis. Les premiers stades de développement. Insectes Soc 2:141–158CrossRefGoogle Scholar
  17. Detrain C, Pasteels JM (1991) Caste differences in behavioral thresholds as a basis for polyethism during food recruitment in the ant, Pheidole pallidula (Nyl) (Hymenoptera, Myrmicinae). J Ins Behav 4:157–176CrossRefGoogle Scholar
  18. Detrain C, Pasteels JM (1992) Caste polyethism and collective defense in the ant, Pheidole pallidula—the outcome of quantitative differences in recruitment. Behav Ecol Sociobiol 29:405–412CrossRefGoogle Scholar
  19. Fernandez-Marin H, Zimmermann JK, Wcislo WT (2003) Nest-founding in Acromyrmex octospinosus (Hymenoptera, Formicidae, Attini): demography and putative prophylactic behaviors. Insectes Soc 50:304–308CrossRefGoogle Scholar
  20. Fewell JH, Page RE (2000) Colony-level selection effects on individual and colony foraging task performance in honeybees, Apis mellifera L. Behav Ecol Sociobiol 48:173–181CrossRefGoogle Scholar
  21. Fjerdingstad EJ, Crozier RH (2006) The evolution of worker caste diversity in social insects. Am Nat 167:390–400PubMedCrossRefGoogle Scholar
  22. Franks NR, Deneubourg J-L (1997) Self-organizing nest construction in ants: individual worker behaviour and the nest’s dynamics. Anim Behav 54:779–796PubMedCrossRefGoogle Scholar
  23. Gautrais J, Theraulaz G, Deneubourg JL, Anderson C (2002) Emergent polyethism as a consequence of increased colony size in insect societies. J Theor Biol 215:363–373PubMedCrossRefGoogle Scholar
  24. Gorelick R, Bertram SM, Killeen PR, Fewell JH (2004) Normalized mutual entropy in biology: quantifying division of labor. Am Nat 164:678–682CrossRefGoogle Scholar
  25. Jeanne RL (1991a) Polyethism. In: Ross KG Matthews RW (eds) The social biology of wasps Cornell University Press Ithaca, New York , pp 389 – 425Google Scholar
  26. Jeanne RL (1991b) The swarm-founding Polistinae. In: Ross KG, Matthews RW (eds) The social biology of wasps. Cornell University Press, Ithaca, New York, pp 191– 231Google Scholar
  27. Jeanson R, Kukuk PF, Fewell JH (2005) Emergence of division of labour in halictine bees: contributions of social interactions and behavioural variance. Anim Behav 70:1183–1193CrossRefGoogle Scholar
  28. Jones JC, Myerscough MR, Graham S, Oldroyd BP (2004) Honey bee nest thermoregulation: diversity promotes stability. Science 305:402–404PubMedCrossRefGoogle Scholar
  29. Karsai I, Wenzel JW (1998) Productivity, individual-level and colony-level flexibility, and organization of work as consequences of colony size. Proc Natl Acad Sci U S A 95:8665–8669PubMedCrossRefGoogle Scholar
  30. Kolmes SA (1985) A quantitative study of the division of labor among worker honey bees. Z Tierpsychol 68:287–302Google Scholar
  31. Kolmes SA, Winston ML (1988) Division of labor among worker honey bees in demographically manipulated colonies. Insectes Soc 35:262–270CrossRefGoogle Scholar
  32. Lachaud J-P, Fresneau D (1987) Social regulation in ponerine ants. In: Deneubourg JL, Pasteels J (eds) From individual to collective behavior in social insects, 54. Birkhäuser-Verlag, Basel, pp 197– 217Google Scholar
  33. Mailleux AC, Deneubourg JL, Detrain C (2003) How does colony growth influence communication in ants? Insectes Soc 50:24–31CrossRefGoogle Scholar
  34. McCarthy MC, Enquist BJ (2005) Organismal size, metabolism and the evolution of complexity in metazoans. Evol Ecol Res 7:681–696Google Scholar
  35. Merkle D, Middendorf M (2004) Dynamic polyethism and competition for tasks in threshold reinforcement models of social insects. Adapt Behav 12:251–262CrossRefGoogle Scholar
  36. Meudec M (1979) Le comportement d’émigration chez la fourmi Tapinoma erraticum (Dolichoderinae): un exemple de régulation sociale. Bull Biol Fr Belg 13:321–374Google Scholar
  37. Michener CD (1974) The social behavior of the bees. A comparative study. Belknap Press of Harvard University Press, Cambridge, MAGoogle Scholar
  38. Murakami T, Higashi S, Windsor D (2000) Mating frequency, colony size, polyethism and sex ratio in fungus-growing ants (Attini). Behav Ecol Sociobiol 48:276–284CrossRefGoogle Scholar
  39. Myerscough MR, Oldroyd BP (2004) Simulation models of the role of genetic variability in social task allocation. Insectes Soc 51:146–152CrossRefGoogle Scholar
  40. Naug D (2001) Ergonomic mechanisms for handling variable amounts of work in colonies of the wasp Ropalidia marginata. Ethology 107:1115–1123CrossRefGoogle Scholar
  41. Naug D, Gadagkar R (1999) Flexible division of labor mediated by social interactions in an insect colony—a simulation model. J Theor Biol 197:123–133PubMedCrossRefGoogle Scholar
  42. O’Donnell S (1995) Division of labor in postemergence colonies of the primitively eusocial wasp Polistes instabilis De Saussure (Hymenoptera, Vespidae). Insectes Soc 42:17–29CrossRefGoogle Scholar
  43. O’Donnell S (1998) Dominance and polyethism in the eusocial wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae). Behav Ecol Sociobiol 43:327–331CrossRefGoogle Scholar
  44. Oldroyd BP, Fewell JH (2007) Genetic diversity promotes homeostasis in insect colonies. Trends Ecol Evol 22:408–413PubMedCrossRefGoogle Scholar
  45. Oster GF, Wilson EO (1978) Caste and Ecology in the Social Insects. Princeton University Press, Princeton, New JerseyGoogle Scholar
  46. Pacala SW, Gordon DM, Godfray HCJ (1996) Effects of social group size on information transfer and task allocation. Evol Ecol 10:127–165CrossRefGoogle Scholar
  47. Page RE, Mitchell SD (1998) Self-organization and the evolution of division of labor. Apidologie 29:171–190CrossRefGoogle Scholar
  48. Pankiw T, Page RE (2000) Response thresholds to sucrose predict foraging division of labor in honeybees. Behav Ecol Sociobiol 47:265–267CrossRefGoogle Scholar
  49. Plowright RC, Plowright CMS (1988) Elitism in social insects: a positive fed-back model. In: Jeanne RL (ed) Interindividual behavioral variability in social insects. Westview Press, Boulder, CO, pp 419– 431Google Scholar
  50. Robinson GE, Page RE (1988) Genetic determination of guarding and undertaking in honey-bee colonies. Nature 333:356–358CrossRefGoogle Scholar
  51. Robinson GE, Page RE (1989) Genetic basis for division of labor in an insect society. In: Breed MD, Page RE (eds) The genetics of social evolution. Westview Press, Boulder, CO, pp 61 – 80Google Scholar
  52. Schatz B (1997) Modalités de recherche et de la récolte alimentaire chez le fourmi Ectatomma ruidum Roger: flexibilités individuelle et collective. PhD Thesis. Université Paul Sabatier, Toulouse, pp 275Google Scholar
  53. Seeley TD (1995) The wisdom of the hive. Harvard University Press, Cambridge, MassachusettsGoogle Scholar
  54. Shannon CE (1948) A mathematical theory of communication. Bell Syst Tech J 27:379–423, 623–656Google Scholar
  55. Stuart RJ, Page RE (1991) Genetic component to division of labor among workers of a Lepthoracine ant. Naturwissenschaften 78:375–377CrossRefGoogle Scholar
  56. Theraulaz G, Bonabeau E, Deneubourg J-L (1998) Response threshold reinforcement and division of labour in insect societies. Proc R Soc Lond B 265:327–332CrossRefGoogle Scholar
  57. Thomas ML, Elgar MA (2003) Colony size affects division of labour in the ponerine ant Rhytidoponera metallica. Naturwissenschaften 90:88–92PubMedGoogle Scholar
  58. Traniello JFA (1978) Caste in a primitive ant: absence of age polyethism in Amblyopone. Science 202:770–772PubMedCrossRefGoogle Scholar
  59. Traniello JFA, Rosengaus RB (1997) Ecology, evolution and division of labour in social insects. Anim Behav 53:209–213CrossRefGoogle Scholar
  60. Waibel M, Floreano D, Magnenat S, Keller L (2006) Division of labour and colony efficiency in social insects: effects of interactions between genetic architecture, colony kin structure and rate of perturbations. Proc R Soc Lond B 273:1815–1823CrossRefGoogle Scholar
  61. Weidenmüller A (2004) The control of nest climate in bumblebee (Bombus terrestris) colonies: interindividual variability and self reinforcement in fanning response. Behav Ecol 15:120–128CrossRefGoogle Scholar
  62. Wilson EO (1976) Behavioral discretization and number of castes in an ant species. Behav Ecol Sociobiol 1:141–154CrossRefGoogle Scholar
  63. Wilson EO (1980) Caste and division of labor in leaf-cutter ants (Hymenoptera, Formicidae, Atta).1. The overall pattern in Atta sexdens. Behav Ecol Sociobiol 7:143–156CrossRefGoogle Scholar
  64. Wilson EO (1986) Caste and division of labor in Erebomyrma, a genus of dimorphic ants (Hymenoptera, Formicidae, Myrmicinae). Insectes Soc 33:59–69CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • Raphaël Jeanson
    • 1
  • Jennifer H. Fewell
    • 2
  • Root Gorelick
    • 3
  • Susan M. Bertram
    • 3
  1. 1.Centre de Recherches sur la Cognition Animale, CNRS UMR 5169Université Paul SabatierToulouseFrance
  2. 2.School of Life SciencesArizona State UniversityTempeUSA
  3. 3.Department of BiologyCarleton UniversityOttawaCanada

Personalised recommendations