Recent evidence from cooperative insect, bird and mammal societies has challenged the assumption that teaching is restricted to humans. However, little is known about the factors affecting the degree to which individuals in such societies contribute to teaching. Here, I examine variation in contributions to teaching in meerkats, where older group members teach pups to handle difficult prey. I show that investment in teaching varies with characteristics of pups, helpers, groups and ecological conditions. Although prior experience in caring for pups did not significantly influence teaching behaviour, younger helpers, which were still investing in growth, contributed less to teaching than older individuals. This suggests that, in common with other cooperative activities, contributions to teaching vary with the costs experienced by individual group members. However, in contrast to other forms of helping in meerkats, I detected no effects of nutritional state on teaching, suggesting that it carries relatively low costs. In species where individuals can potentially gain direct or indirect fitness benefits from facilitating learning in others, low costs divided among multiple group members may help tip the balance towards selection for teaching.

References

Barnard, J. A. 2000 Costs and benefits of group foraging in cooperatively breeding meerkats. PhD thesis, University of Cambridge, UK. Google Scholar
Bergmüller R, Johnstone R.A, Russell A.F& Bshary R . 2007 Integrating cooperative breeding into theoretical concepts of cooperation. Behav. Process. 76, 61–72.doi:10.1016/j.beproc.2007.07.001. . Crossref, PubMed, Web of Science, Google Scholar
Boland C.R.J, Heinsohn R& Cockburn A . 1997 Experimental manipulation of brood reduction and parental care in cooperatively breeding white-winged choughs. J. Anim. Ecol. 66, 683–691.doi:10.2307/5921. . Crossref, Web of Science, Google Scholar
Bourke A.F.G Sociality and kin selection in insects. Behavioural ecology: an evolutionary approach , Krebs J.R& Davies N.B . 1997pp. 203–227. Eds. Oxford, UK:Blackwell Science. Google Scholar
Brotherton P.N.M, Clutton-Brock T.H, O'Riain M.J, Gaynor D, Sharpe L, Kansky R& McIlrath G.M . 2001 Offspring food allocation by parents and helpers in a cooperative mammal. Behav. Ecol. 12, 590–599.doi:10.1093/beheco/12.5.590. . Crossref, Web of Science, Google Scholar
Brown J.L Helping and communal breeding in birds. 1987 Princeton, NJ:Princeton University Press. Google Scholar
Caro T.M Cheetahs of the Serengeti plains: grouping in an asocial species. 1994 Chicago, IL:University of Chicago Press. Google Scholar
Caro T.M& Hauser M.D . 1992 Is there teaching in nonhuman animals?. Q. Rev. Biol. 67, 151–174.doi:10.1086/417553. . Crossref, PubMed, Web of Science, Google Scholar
Clutton-Brock T.H . 2002 Breeding together: kin selection and mutualism in cooperative vertebrates. Science. 296, 69–72.doi:10.1126/science.296.5565.69. . Crossref, PubMed, Web of Science, Google Scholar
Clutton-Brock T.H, et al. 1998 Costs of cooperative behaviour in suricates (Suricata suricatta). Proc. R. Soc. B. 265, 185–190.doi:10.1098/rspb.1998.0281. . Link, Web of Science, Google Scholar
Clutton-Brock T.H, O'Riain M.J, Brotherton P.N.M, Gaynor D, Kansky R, Griffin A.S& Manser M . 1999 Selfish sentinels in cooperative mammals. Science. 284, 1640–1644.doi:10.1126/science.284.5420.1640. . Crossref, PubMed, Web of Science, Google Scholar
Clutton-Brock T.H, Brotherton P.N.M, O'Riain M.J, Griffin A.S, Gaynor D, Sharpe L, Kansky R, Manser M.B& McIlrath G.M . 2000 Individual contributions to babysitting in a cooperative mongoose, Suricata suricatta. Proc. R. Soc. B. 267, 301–305.doi:10.1098/rspb.2000.1000. . Link, Web of Science, Google Scholar
Clutton-Brock T.H, Brotherton P.N.M, O'Riain M.J, Griffin A.S, Gaynor D, Kansky R, Sharpe L& McIlrath G.M Contributions to cooperative rearing in meerkats. Anim. Behav. 61, 2001a 705–710.doi:10.1006/anbe.2000.1631. . Crossref, Web of Science, Google Scholar
Clutton-Brock T.H, et al. Cooperation, control, and concession in meerkat groups. Science. 291, 2001b 478–481.doi:10.1126/science.291.5503.478. . Crossref, PubMed, Web of Science, Google Scholar
Clutton-Brock T.H, Russell A.F, Sharpe L.L, Young A.J, Balmforth Z& McIlrath G.M . 2002 Evolution and development of sex differences in cooperative behavior in meerkats. Science. 297, 253–256.doi:10.1126/science.1071412. . Crossref, PubMed, Web of Science, Google Scholar
Cockburn A . 1998 Evolution of helping behavior in cooperatively breeding birds. Annu. Rev. Ecol. Syst. 29, 141–177.doi:10.1146/annurev.ecolsys.29.1.141. . Crossref, Google Scholar
Doolan S.P& Macdonald D.W . 1996 Diet and foraging behaviour of group-living meerkats, Suricata suricatta, in the southern Kalahari. J. Zool. 239, 697–716. Crossref, Web of Science, Google Scholar
Eden S.F . 1987 When do helpers help? Food availability and helping in the moorhen, Gallinula chloropus. Behav. Ecol. Sociobiol. 21, 191–195.doi:10.1007/BF00303210. . Crossref, Web of Science, Google Scholar
Emlen S.T Evolution of cooperative breeding in birds and mammals. Behavioural ecology: an evolutionary approach , Krebs J.R& Davies N.B . 1991pp. 301–337. Eds. Oxford, UK:Blackwell Science. Google Scholar
Franks N.R& Richardson T . 2006 Teaching in tandem-running ants. Nature. 439, 153 doi:10.1038/439153a. . Crossref, PubMed, Web of Science, Google Scholar
Gergely G, Egyed K& Király I . 2007 On pedagogy. Dev. Sci. 10, 139–146.doi:10.1111/j.1467-7687.2007.00576.x. . Crossref, PubMed, Web of Science, Google Scholar
Heinsohn R.G . 1991 Slow learning of foraging skills and extended parental care in cooperatively breeding white-winged choughs. Am. Nat. 137, 864–881.doi:10.1086/285198. . Crossref, Web of Science, Google Scholar
Heinsohn R& Legge S . 1999 The cost of helping. Trends Ecol. Evol. 14, 53–57.doi:10.1016/S0169-5347(98)01545-6. . Crossref, PubMed, Web of Science, Google Scholar
Jacobs D.S& Jarvis J.U.M . 1996 No evidence for the work-conflict hypothesis in the eusocial naked mole-rat (Heterocephalus glaber). Behav. Ecol. Sociobiol. 39, 401–409.doi:10.1007/s002650050307. . Crossref, Web of Science, Google Scholar
Jordan N.R, Cherry M.I& Manser M.B . 2007 Latrine distribution and patterns of use by wild meerkats: implications for territory and mate defence. Anim. Behav. 73, 613–622.doi:10.1016/j.anbehav.2006.06.010. . Crossref, Web of Science, Google Scholar
Kokko H, Johnstone R.A& Clutton-Brock T.H . 2001 The evolution of cooperative breeding through group augmentation. Proc. R. Soc. B. 268, 187–196.doi:10.1098/rspb.2000.1349. . Link, Web of Science, Google Scholar
Komdeur J . 1994 The effect of kinship on helping in the cooperative breeding Seychelles warbler, Acrocephalus seychellensis. Proc. R. Soc. B. 256, 47–52.doi:10.1098/rspb.1994.0047. . Link, Web of Science, Google Scholar
Komdeur J . 1996 Influence of helping and breeding experience on reproductive performance in the Seychelles warbler: a translocation experiment. Behav. Ecol. 7, 326–333.doi:10.1093/beheco/7.4.417. . Crossref, Web of Science, Google Scholar
Margulis S.W, Nabong M, Alaks G, Walsh A& Lacy R.C . 2005 Effects of early experience on subsequent parental behaviour and reproductive success in oldfield mice, Peromyscus poliolotus. Anim. Behav. 69, 627–634.doi:10.1016/j.anbehav.2004.04.021. . Crossref, Web of Science, Google Scholar
Premack D& Premack A.J Why animals lack pedagogy and some cultures have more of it than others. Handbook of education and human development: new models of learning, teaching and schooling , Olson D.R& Torrance N . 1996pp. 302–323. Eds. Cambridge, MA:Blackwell. Google Scholar
Raihani N.J& Ridley A.R . 2008 Experimental evidence for teaching in wild pied babblers. Anim. Behav. 75, 3–11.doi:10.1016/j.anbehav.2007.07.024. . Crossref, Web of Science, Google Scholar
Richardson T.O, Sleeman P.A, McNamara J.M, Houston A.I& Franks N.R . 2007 Teaching with evaluation in ants. Curr. Biol. 17, 1520–1526.doi:10.1016/j.cub.2007.08.032. . Crossref, PubMed, Web of Science, Google Scholar
Russell A.F, Sharpe L.L, Brotherton P.N.M& Clutton-Brock T.H . 2003 Cost minimization by helpers in cooperative vertebrates. Proc. Natl Acad. Sci. USA. 100, 3333–3338.doi:10.1073/pnas.0636503100. . Crossref, PubMed, Web of Science, Google Scholar
Schall R . 1991 Estimation in generalized linear models with random effects. Biometrika. 78, 719–727.doi:10.1093/biomet/78.4.719. . Crossref, Web of Science, Google Scholar
Tardif S.D, Richter C.B& Carson R.L . 1984 Effects of sibling-rearing experience on future reproductive success in 2 species of Callitrichidae. Am. J. Primatol. 6, 377–380.doi:10.1002/ajp.1350060408. . Crossref, PubMed, Web of Science, Google Scholar
Thornton A& McAuliffe K . 2006 Teaching in wild meerkats. Science. 313, 227–229.doi:10.1126/science.1128727. . Crossref, PubMed, Web of Science, Google Scholar
Thornton, A. & Raihani, N. J. 2008 The evolution of teaching. Anim. Behav. (doi:10.1016/j.anbehav.2007.12.014). Google Scholar
West S.A, Griffin A.S& Gardner A . 2007 Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection. J. Evol. Biol. 20, 415–432.doi:10.1111/j.1420-9101.2006.01258.x. . Crossref, PubMed, Web of Science, Google Scholar