Readings
Topics for Discussions:
1.
The
problem of semantics: what does cooperation mean? What does altruism mean?
2.
Social
evolution theory: why does helping evolve?
3.
Game
theory: what helping strategies lead to optimal outcomes in games?
4.
Why
are some animals cooperative breeders?
5.
What
is the relative importance of direct vs indirect
benefits of helping?
6.
What
prevents cheating in mutualisms between species?
7. What are the benefits of social bonds?
8.
How
does partner choice and partner switching drive helping?
9. Does reciprocity exist in nature outside humans?
10.What are the cognitive
requirements for reciprocity?
11. Are animals capable of reciprocity?
12. How do organisms recognize kin?
13. Do nonhuman animals possess psychological
altruism?
14. Evolution of cooperation in humans
15. What prevents cheating during bacteria
and slime mold cooperation?
16. Why are some animals
eusocial?
17. When is helping not cooperation?
Semantics: What are we talking about?
What does altruism mean?
What does cooperation mean?
Required:
West
et al. (2007) Social semantics: altruism, cooperation, mutualism, strong reciprocity
and group selection. J Evol. Biol. 20:415-32
West
et al. (2008) Social
semantics: how useful has group selection been? J Evol. Biol.
Optional:
Connor, R.C.
1986. Pseudo-reciprocity – investing in mutualism. Anim. Behav. 34: 1562–1566.
Connor, R.C.
1995. Altruism among nonrelatives – alternatives to the
prisoners-dilemma. Trends Ecol. Evol. 10: 84–86.
Leimar, O. &
Connor, R. C. in Genetic and Cultural Evolution of Cooperation (ed.
Hammerstein, P.) 202–222 (MIT Press, 2003).
Lehmann, L. & Keller, L. (2006).
The evolution of cooperation and altruism. A general framework and classification
of models. Journal of Evolutionary Biology, 19, 1365-1378.
Lehmann, L., Keller, L., West, S. A.
& Roze, D. (2007). Group selection and kin
selection. Two concepts but one process. Proc. Natl. Acad. Sci. U.S.A., 104,
6736-6739.
Leigh. 2010. The group selection controversy. J Evol Biol.
Social Evolution Theory: inclusive
fitness and alternative approaches
Why does helping evolve?
Required:
Maynard Smith, J. (1964). Group selection and kin selection. Nature, 201, 1145-1147.
West et al. (2002) Cooperation and competition between relatives. Science 296, 72-75.
Gardner, A. (2008). Primer: The Price equation. Current Biology, 18, R198-R202.
Optional:
Hamilton, W. D. (1963). The
evolution of altruistic behaviour. Am. Nat., 97,
354-356.
Hamilton, W. D. (1964). The genetical evolution of social behaviour,
I & II. J. Theor. Biol
Michod,
R. E. & Hamilton, W. D. (1980). Coefficients of relatedness in
sociobiology. Nature, 288, 694-697.
Grafen, A. (1982). How not to measure inclusive
fitness. Nature, 298, 425-426.
Griffin
and West. Kin selection: fact and fiction. Trends in Ecology & Evolution
(2002) vol. 17 (1) pp. 15-21.
Frank, S. A. (2003). Repression of
competition and the evolution of cooperation. Evolution, 57,
693-705.
Gardner,
A., West, S. A. & Barton, N. H. (2007). The relation between multilocus population genetics and social evolution theory.
American Naturalist, 169, 207-226.
Grafen, A. (2007). The formal Darwinism project:
a mid-term report. Journal of Evolutionary Biology, 20,
1243-1254.
Grafen, A. (2007). Detecting kin selection at work using inclusive fitness. Proc. Roy. Soc. Lond. B, 274, 713-719.
Lehmann, L., Keller, L. & Sumpter, D. J. T. (2007). Inclusive fitness explains the
evolution of helping and harming behaviors on graphs. Journal of
Evolutionary Biology, 20, 2284-2295.
West
et al. Evolutionary Explanations for Cooperation. Current Biology (2007) vol.
17 (16) pp. R661-R672
West, S.A., El
Mouden, C. & Gardner, A. In press. Social evolution theory and its application to the evolution of
cooperation in humans.
Game theory: Frequency-dependent
selection in a simplified world
What helping strategies
lead to optimal outcomes in games?
Required:
Axelrod
and Hamilton. (1981) The evolution of cooperation. Science 211:1390-1397
Optional:
Boyd, R. & Richerson, P. J. 1988 The evolution of reciprocity in sizable groups. J. Theor. Biol. 132, 337–356.
Bshary, R. & Bronstein, J.L. (2004) Game structures in mutualistic interactions: what can the evidence tell us about the kind of models we need? Adv. Study Behav. 34: 59–101.
Bshary, R. & Bergmuller. (2008) Distinguishing four fundamental approaches to the evolution of helping. J. Evol Biol 21: 405-420
Mcnamara et al. (2008) The coevolution of choosiness and cooperation. Nature 451:189-192
Nowak MA, K Sigmund (1998). Evolution of indirect reciprocity by image scoring. Nature 393: 573-577.
Nowak MA, K Sigmund (2004). Evolutionary dynamics of biological games. Science 303: 793-799.
Nowak MA, K Sigmund (2005). Evolution of indirect reciprocity. Nature 437: 1291-1298.
Nowak MA (2006). Five rules for the evolution of cooperation. Science 314: 1560-1563.
Pfeiffer et al.(2005) Evolution of cooperation by generalized reciprocity. Proceedings of the Royal Society B: Biological Sciences 272: 1115-1120
Cooperative breeding and sociality in
birds and mammals
Why are some animals
cooperative breeders?
Required:
Bergmuller et al. (2007) Integrating
cooperative breeding into theoretical concepts of cooperation. Behavioural Processes 76: 61-72
Optional:
Bergmuller, Heg, and Taborksy. (2005) Helpers in a cooperatively breeding cichlid stay and pay or disperse and breed, depending on ecological constraints. Proceedings Of The Royal Society B-Biological Sciences 272:325-331.
Bergmuller et al. (2007) On the further integration of cooperative breeding and cooperation theory. Behavioural Processes 76: 170-181
Bruintjes and Taborsky. (2008) Helpers in a cooperative breeder pay a high price to stay: effects of demand, helper size and sex. Animal behaviour 75: 1843-1850
Canestrari, Vera, Chiarati, Marcos,Vila and Baglione. (2010) False feeding: the trade-off between chick hunger and caregivers needs in cooperative crows. Behavioral Ecology 21:1-9.
Doerr, Doerr, and Safran. (2007) Integrating delayed dispersal into broader concepts of social group formation. Behavioural Processes 76: 114-117.
Emlen, S.T., Reeve, H.K., & Keller, L. (1998) Reproductive skew: disentangling concessions from control. Trends in Ecology and Evolution, 13: 458-459.
Jennions, M. D. & Macdonald, D. W. (1994). Cooperative breeding in mammals. Trends Ecol. Evol., 9, 89-93.
Kokko, H., Johnstone, R. A. & Clutton-Brock, T. H. (2001). The evolution of cooperative breeding through group augmentation. Proc. R. Soc. Lond. B, 268, 187-196.
McDonald, Kazem, Clarke, and Wright. (2008) Helping as a signal: does removal of potential audiences alter helper behavior in the bell miner? Behavioral Ecology 19:1047-1055.
Richardson, Burke, and Komdeur. (2002) Direct benefits and the evolution of female-biased cooperative breeding in Seychelles warblers. Evolution 56: 2313-2321.
Safran, Doerr, Sherman, Doerr, Flaxman, and Winkler. (2007) Group breeding in vertebrates: linking individual and population-level approaches. Evolutionary Ecology Research 9: 1163-1185.
Taborsky (Barbara),
Skubic, and Bruintjes. (2007) Mothers adjust egg size to helper number
in a cooperatively breeding cichlid. Behavioral Ecology 18:
652-657.
Wiley,
R. H. & Rabenold, K. N. (1984). The evolution of
cooperative breeding by delayed reciprocity and queuing for favorable social
positions. Evolution, 38, 609-621.
What ecological conditions
lead to cooperative breeding?
Rubenstein, D.R. and I.J. Lovette.
2007. Temporal environmental variability drives the evolution of cooperative
breeding in birds. Current Biology
17:1414-1419.
Lion
and Gandon. Habitat saturation and the spatial
evolutionary ecology of altruism. Journal of evolutionary biology (2009) vol.
22 (7) pp. 1487-1502
Gardner
and Foster. The Evolution and Ecology of Cooperation–History and
Concepts. Ecology of Social Evolution. Berlin, Heidelberg: Springer (2008)
What is the relative
importance of direct vs indirect benefits of helping?
Required:
Clutton-Brock, T.H. (2009) Cooperation between non-kin in
animal societies. Nature 461: 51-57
Schino and Aureli.
(2010) The relative roles of kinship and reciprocity in explaining primate
altruism. Ecology Letters. 12:1-6.
Optional:
Clutton-Brock, T. H. (2002). Breeding together: kin selection, reciprocity and mutualism in cooperative animal societies. Science, 296, 69-72.
Cornwallis, C., West, S. and Griffin, A. (in press) Routes to indirect fitness in cooperatively breeding vertebrates: kin discrimination and limited dispersal.
Emlen, S.T.& Wrege,P.H. The role of kinship in
helping decisions among white-fronted bee-eaters. Behav.
Ecol. Sociobiol. 23, 305–315 (1988).
Griffin
and West. Kin discrimination and the benefit of helping in cooperatively
breeding vertebrates. Science (2003) vol. 302 (5645) pp. 634
Russell,
A. F. & Hatchwell, B. J. (2001). Experimental
evidence for kin-biased helping in
Sherman, P. W. (1977). Nepotism and the evolution of alarm calls. Science, 197, 1246-1253.
Viblanc et al. (2009) Kin selection in Columbian ground squirrels (Urocitellus columbianus): littermate kin provide individual fitness benefits. Proceedings of the Royal Society B: Biological Sciences 1-7.
Mutualisms between species: What prevents cheating between species?
Strassmann,
J.E. and Queller, D.C. (2010) The social organism:
congresses, parties and committees. Evolution 64:605-616.
Jander, K. C. and
Herre, E.A. (2010) Host sanctions and pollinator cheating
in the fig tree-fig wasp mutualism. Proc. Roy. Soc. Lond. B published online Jan 13.
Raihani NJ, Grutter AS, Bshary R (2010) Punishers benefit from third-party punishment in fish. Science 327: 171-171.
Fig and fig-wasp mutualism
Cleaner and client fish mutualism
Bshary,
R. (2002). Biting cleaner fish use altruism to deceive image-scoring client
reef fish Proc. Roy. Soc. Lond. B, 269,
2087-2093.
Bshary,
R. & SchŠffer, D. (2002). Choosy reef fish select
cleaner fish that provide high- quality service. Animal Behaviour,
63, 557-564.
Bshary,
R. & Grutter, A. S. (2002). Asymetric
cheating opportunities and partner control in a cleaner fish mutualism. Animal
Behaviour, 63, 547-555.
What are the benefits of social bonds?
Required:
Optional:
Aureli,F.,van Schaik,C.P. & van Hoof,J. (1989) Functional aspects of reconciliation among captive long-tailed macaques. Am.
J. Primatol. 19, 39–51.
Aureli, F.
(1992) Post-conflict behaviour among wild long-tailed
macaques Macaca fascicularis.
Behav. Ecol. Sociobiol. 31,
329–337
Boogert NJ, Reader SM, Laland KN (2006) The relation between social
rank, neophobia and individual learning in starlings. Anim. Behav. 72:
1229-1239.
deWaal,F.B.M.The integration of dominance and social bonding in primates. Q. Rev. Biol. 61, 459–479 (1986).
Reyes, T.L. & Mateo, J.M. 2008. Oxytocin and cooperation: cooperation with non-kin associated with mechanisms for affiliation. The Journal of Social, Evolutionary & Cultural Psychology, 2, 234-246.
Seed, A. M., Clayton, N. S. & Emery, N. J. 2008 Cooperative problem solving in rooks (Corvus frugilegus). Proc. R. Soc. B 2008 275, 1421-1429
Silk, JB. 2007. Social components of fitness in primate groups. Science 317:1347-1351.
Silk,
J.B., Altmann, J., & Alberts, S.C. 2006. Social relationships among
adult female baboons (Papio cynocephalus) I. Variation in the strength
of social bonds. Behavioral Ecology and Sociobiology 61: 183-195
Silk, J.B., Alberts, S.C., Altmann, J.
2006. Social Relationships among Adult Female Baboons (Papio
cynocephalus) II: Variation in the Quality and Stability of Social
Bonds. Behavioral Ecology and Sociobiology 61:197-204
Weidt et al. (2008) Not only mate choice matters: fitness consequences of social partner choice in female house mice. Animal behaviour 75: 801-808
How do individuals punish?
Mulder, R. A. & Langmore,
N. E. (1993). Dominant males punish helpers for temporary defection in superb
fairy wrens. Animal Behaviour, 45,
830-833.
How does partner choice and
partner switching drive helping?
Required:
Optional:
de Waal FBM (1997) The chimpanzees service economy - food for grooming. Evol Hum Behav 18:375–386.
Gumert, MD (2007) Payment for sex in a macaque mating market. Animal Behaviour 74:1655-1667.
Noe, R. & Hammerstein, P. (1995) Biological markets. Trends in Ecology & Evolution 10:336-339.
Noe, R. (2001) Biological markets: partner choice as the driving force behind the evolution of cooperation. In: Economics in
Nature (R. Noe , J. A. R. A. M. van Hooff & P. Hammerstein, eds), pp. 93–118. Cambridge University Press, Cambridge.
Schwartz,
M.W. & Hoeksema, J.D. 1998. Specialization and resource trade:
Biological markets as a model of mutualisms. Ecology 79:
1029–1038.
Reciprocity
Does reciprocity exist in
nature outside humans?
Required:
de Waal, F. B. M. & Brosnan, S. F. (2006). Simple and complex reciprocity in primates. In: Cooperation in primates and humans: mechanisms and evolution. (eds. P.M. Kappeler & Carel P. van Schaik) pp. 85-105, Springer, New York.
Optional:
Brosnan, S. F., & de Waal, F. B. M. (2002). A proximate perspective on reciprocal altruism. Human Nature 13: 129-152.
Hart, B. L. and L. A. Hart (1992). Reciprocal allogrooming in impala, Aepyceros melampus. Anim. Behav. 44: 1073-1083.
Packer, C. (1977) Reciprocal altruism in olive baboons (Papio anubis). Nature 265: 441–443.
Seyfarth, R. M. and D. L. Cheney (1984). Grooming, Alliances and Reciprocal Altruism in Vervet Monkeys. Nature 308: 541-543.
Wilkinson, G.S. (1984) Reciprocal food sharing in the vampire bat. Nature 308: 181-184.
Wilkinson,
G. S. (1988) Reciprocal altruism in bats and other mammals. Ethology and Sociobiology (1988) 9: 85-100.
What are the cognitive
requirements for reciprocity?
Required:
Optional:
LEE D (2008) Game theory and neural basis of social decision making. NATURE NEUROSCIENCE 11:404-409
Ramseyer et al. (2005) Accepting loss: the temporal limits of reciprocity in brown capuchin monkeys. Proceedings of the Royal Society B: Biological Sciences 273: 179-184
Rilling, J. K., D. A. Gutman, et al. (2002). A neural basis for social cooperation. Neuron 35: 395-405.
Rilling JK, King-Casas B, Sanfey AG (2008) The neurobiology
of social decision-making. CURRENT OPINION IN
NEUROBIOLOGY 18: 159-165.
Schultz, W. 2004 Neural coding of basic reward terms of animal learning theory, game theory, microeconomics and behavioural ecology. CURRENT OPINION IN NEUROBIOLOGY 14: 139-147
Stephens DW, McLinn CM, Stevens JR (2006) Effects of temporal clumping and payoff accumulation on impulsiveness and cooperation. BEHAVIOURAL PROCESSES 71: 29-40.
Stevens, J. R., Cushman, F. A. & Hauser, M. D. (2005). Evolving the psychological mechanisms for cooperation. Annu. Rev. Ecol. Syst., 36, 499-518.
Yamamoto S, Tanaka M (2009) Do chimpanzees (Pan troglodytes) spontaneously take turns in a reciprocal cooperation task? J Comp Psychol 123: 242–249
Yamamoto S, Tanaka M
(2009) Development of selfish tactics but not reciprocal cooperation in
chimpanzee (Pan troglodytes) mother-infant pairs in social problem
situations. Anim Cogn 12: 27–36.
What is required to demonstrate reciprocity?
Required:
Optional:
Brosnan SF, Silk JB, Henrich J, et al. (2009) Chimpanzees (Pan troglodytes) do
not develop contingent reciprocity in an experimental task. Animal Cognition 12: 587-597
Clements, KC, Stephens, DW (1995) Testing models of nonkin cooperation - Mutualism and the Prisoner's Dilemma. Animal Behaviour 50: 527-535
Hauser,
M. D., M. K. Chen, et al. (2003). Give unto others: genetically
unrelated cotton-top tamarin monkeys preferentially give food to those
who altruistically give food back. Proceedings of the Royal Society of
London Series B-Biological Sciences 270: 2363-2370
Krams I, Berzins A, Krama T, et al. (2010) The increased risk of predation enhances cooperation. Proceedings of the Royal Society of
London Series B-Biological Sciences 277: 513-518.
Melis, A. P., B. Hare, et al. (2008). Do chimpanzees reciprocate received favours? Animal Behaviour 76: 951-962. VIDEO doi:10.1016/j.anbehav.2008.05.014
Noe, R. (2006) Cooperation experiments: coordination through communication versus acting apart together. Animal Behaviour 71: 1-18. doi:10.1016/j.anbehav.2005.03.037
Rutte, C. and Taborsky, M. (2007)
Generalized Reciprocity in Rats. PLoS Biology 5:e196
How do birds and mammals
recognize kin?
Required:
Optional:
El-Showk, S. et al. (2009) Are you my mother? Kin recognition in the ant Formica fusca. J. Evol. Biol. 23-397-406.
Gardner,
A. and S.A. West
(2009) Greenbeards.
Evolution 64:25-38.
Grafen, A. (1990) Do animals really recognise kin? Anim. Behav., 39,
42-54.
Hauber, M. E. & Sherman, P. W. (2001)
Self-referent phenotype matching: theoretical considerations and empirical
evidence. Trends in Neurosciences, 24, 609-616.
Pfennig, D. W., Collins, J. P. & Ziemba, R. E. (1999) A test of alternative hypotheses for kin recognition in cannibalistic tiger salamanders. Behav. Ecol., 10, 436-443.
Raby CR, Clayton NS (2009) Prospective cognition in animals Behav Proc. 80: 314-324
Do nonhuman animals possess
psychological altruism?
Required:
Grenier, F, Luthi A (2010) Mouse brains wired for empathy? Nature Neuroscience 13: 406-408.
Optional:
Chen QL et al. (2009) Empathy Is Moderated by Genetic Background in Mice. PLOS ONE 4: e4387.
Pokorny JJ, de Waal FBM (2009) Monkeys recognize the faces of group mates in photographs. Proc. Natl. Acad. Sci., USA 106: 21539-21543.
Wynne, CDL (2010) The limits of empathy. Ethology 116:383-384.
Evolution of cooperation in humans
Required:
Boyd R & Richerson PJ (2009) Culture and the evolution of human cooperation. Phil Trans R Soc B 364: 3281-3288.
Andre, J.B. (2010) The evolution of reciprocity: Social types or social incentives? American Naturalist 175: 197-210
Bowles, S. (2009) Did Warfare Among Ancestral Hunter-Gatherers Affect the Evolution of Human Social Behaviors? Science 324, 1293-1298
Boyd, R., H. Gintis, and S. Bowles (2010) Coordinated Punishment of Defectors Sustains Cooperation and Can Proliferate When Rare. Science 328, 617-620
Efferson,C., R. Lalive, and E. Fehr (2008) The Coevolution of Cultural Groups and Ingroup Favoritism. Science 321, 1844-1849
Gachter, S. and B. Herrmann (2009) Reciprocity, culture and human cooperation: previous insights and a new cross-cultural experiment. Phil Trans R Soc B 364, 791-806
Kosfeld, M., Heinrichs, M., Zak, P.J., Fischbacher, U. & Fehr, E. (2005) Oxytocin increases trust in humans. Nature 435: 673–676.
Legume-rhizobia symbiosis
Quorum-sensing and the production of public goods in microorganisms
Buttery
et al. 2009. Quantification of Social Behavior in D. discoideum
Reveals Complex Fixed and Facultative Strategies. Current Biology, Volume 19, Issue 16, Pages 1373-1377.
Diggle, S. P., Griffin, A. S., Campbell, G. S. & West, S. A. (2007). Cooperation and conflict in quorum-sensing bacterial populations. Nature, 450, 411-414.
Gilbert, O.M., K.R. Foster, N.J. Mehdiabadi, J.E. Strassmann and D.C. Queller, (2007) High relatedness maintains multicellular cooperation in a social amoeba by controlling cheater mutants, Proc. Natl. Acad. Sci. USA 104: 8913–8917
Griffin, A. S., West, S. A. & Buckling, A. (2004). Cooperation and competition in pathogenic bacteria. Nature, 430, 1024-1027.
Harcombe, W. 2010 Novel cooperation experimentally evolved between species. Evolution
Kummerli, R., Griffin, A.S., West, S.A., Buckling, A. & Harrison, F. (2009) Viscous medium promotes cooperation in the pathogenic bacterium Pseudomonas aeruginosa. Proceedings of the Royal Society London Series B 276, 3531-3538.
Ross-Gillespie et al. Frequency dependence and cooperation: theory and a test with bacteria. The American Naturalist (2007) vol. 170 (3) pp. 331-342
Queller, D. C. (2004). Kinship is relative. Nature,
430, 975-976.
West, S. A., Griffin, A. S., Gardner, A. & Diggle, S. P. (2006a). Social evolution theory for microbes. Nature Reviews Microbiology, 4, 597-607.
What prevents cheating in diffuse mutualisms, e.g. seed dispersal?
Tiffany, B.H. (2004). Vertebrate dispersal of seed plants through time. Ann. Rev. Ecol. Syst. 35, 1-29.
Vander
Wall, S.B., (2003) Effects of seed-size of wind-dispersed pines (Pinus)
on secondary seed dispersal and the caching behavior of rodents. Oikos,
82, 25-34.
Vander
Wall, S.B., Longland, W.S. (2004) Diplochory: are two seed dispersers
better than one? Trends in Ecology and Evolution, 19, 155-161.
Vander Wall, S.B., Kuhn, K.M., Beck, M.J. (2005) Seed removal, seed predation, and secondary dispersal. Ecology, 86, 801-806
Eusociality
Why are some animals eusocial?
Required:
Hughes,
W. O. H., Oldroyd, B. P., Beekman,
M. & Ratnieks, F. L. W. (2008). Ancestral
monogamy shows kin selection is the key to the evolution of eusociality.
Science, 320, 1213-1216.
Gardner,
A. & Grafen, A. (2009). Capturing the superorganism: a formal theory of group adaptation. Journal
of Evolutionary Biology, 22, 659-671.
Optional:
Frank,
S. A. (1995b). Mutual policing and repression of competition in the evolution
of cooperative groups. Nature, 377, 520-522.
Bernasconi,
G. & Strassman, J. E. (1999). Cooperation among
unrelated individuals: the ant foundress case. Trends
Ecol. Evol., 14, 477-482.
Queller, D. C., Zacchi,
F., Cervo, R., Turillazzi,
S., Henshaw, M. T., Santorelli,
L. A. & Strassmann, J. E. (2000). Unrelated
helpers in a social insect. Nature, 405, 784-787.
Wilson,
E. O. & Hšlldobler, B. (2005). Eusociality: origin and consequences. Proc. Natl. Acad.
Sci. U.S.A., 102, 13367-13371.
Wenseleers, T. & Ratnieks,
F. L. W. (2006). Enforced altruism in insect societies. Nature, 444,
50.
Foster,
K. R., Wenseleers, T. & Ratnieks,
F. L. W. (2006). Kin selection is the key to altruism.
Trends Ecol. Evol., 21, 57-60.
Reeve,
H. K. & Hšlldobler, B. (2007). The emergence of a
superorganism through intergroup competition. Proceedings
of the National Academy of Sciences of the United States of America, 104,
9736-9740.
Burland T M, Bennett N C, Jarvis J U M, Faulkes C G. 2002. Eusociality in
African mole‐rats: new insights from patterns of
genetic relatedness in the Damaraland mole‐rat. Proceedings of the Royal Society of London Series B 269:1025–1030.
Helping behaviours
as non-adaptive byproducts of selected behaviours
When is helping
not cooperation?
Required:
Emlen, S.T., F.L.W. Ratnieks,
H.K. Reeve, J. Shellman-Reeve, P.W. Sherman and P.H. Wrege 1991. Adaptive versus nonadaptive
explanations of behavior: the case of alloparental
helping. American Naturalist, 138:259-270
Optional:
Jamieson.
1991. The Unselected Hypothesis for the Evolution of Helping behaviour.
McCracken.
Communal Nursing in Mexican Free-tailed bats
Tebbich
et al. Social manipulation causes cooperation in keas. Animal Behaviour (1996) vol. 52 (1) pp. 1-10
Bateson, M., Nettle, D. & Roberts, G. (2006). Cues of
being watched enhance