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35.
Samir Okasha, “Why Won’t the Group Selection Controversy Go Away?”
British Journal of the Philosophy of Science
52 (2001), 25–50; Ayelet Shavit, “Shifting Values Partly Explain the Debate Over Group Selection,”
Studies in History and Philosophy of Biological and Biomedical Sciences
35 (2004), 697–720, and
One for All? Facts and Values in the Debates Over Group Selection
(Jerusalem: Magnes Press, 2008) (in Hebrew).

36.
David Sloan Wilson and Edward O. Wilson, “Rethinking the Theoretical Foundations of Sociobiology,”
Quarterly Review of Biology
82 (2007), 327–48. See also Bert H
lldobler and E.O. Wilson,
The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies
(New York: W. W. Norton, 2009). Wilson, in fact, was never as opposed to group selection as either Dawkins or Williams—see Segerstr
le,
Defenders of the Truth
, 37–38. H
lldobler, on the other hand, is not as convinced as Wilson of the importance of group selection. Dawkins, too, has yet to be won over, and the debate remains acrimonious. See his “The Group Delusion,” from January 10, 2009, at RichardDawkins.net. Also, see Ayelet Shavit and Roberta L. Millstein, “Group Selection is Dead! Long Live Group Selection?”
BioScience
58 (2008), 574–75. For an interesting argument for group selection based on the plasticity of circadian clocks in bees, see Guy Bloch, “Plasticity in the Circadian Clock and the Temporal Organization of Insect Societies,” in
Organization of Insect Societies
, ed. J
rgen Gadau and Jennifer Fewell (Cambridge, MA: Harvard University Press, 2009), 402–31.

37.
Each differs in perspective. See S. A. Frank, “George Price’s Contributions to Evolutionary Genetics,”
Journal of Theoretical Biology
175 (1995), 373–88, and “The Price Equation, Fisher’s Fundamental Theorem, Kin Selection, and Causal Analysis,”
Evolution
51 (1997), 1712–29; Okasha, “Why Won’t the Group Selection Controversy Go Away?” A. Grafen, “Developments of the Price Equation and Natural Selection Under Uncertainty,”
Proceedings of the Royal Society London B
, 267 (2000), 1223–27, and “The First Formal Link Between the Price Equation and an Optimization Program,”
Journal of Theoretical Biology
217 (2002), 75–91; but also see I. L. Heisler and J. Damuth, “A Method for Analyzing Selection in Hierarchically Structured Populations,”
American Naturalist
130 (1987), 582–602, which provides a “contextualized” alternative to the Price equation.

38.
Benjamin Kerr and Peter Godfrey-Smith, “Generalization of the Price Equation for Evolutionary Change,”
Evolution
63, no. 2 (2009), 531–36.

39.
The Price equation assumes fixed heritability, and is entirely devoid of mechanism. Though Frank decomposes it neatly, others are concerned that this precludes the interactivity inherent in nature, rendering it good exclusively for border cases. See Okasha, “Why Won’t the Group Selection Controversy Go Away?” for a thorough discussion of the relative merits of the equation. Also Massimo Pigliucci and Jonathan Kaplan,
Making Sense of Evolution: The Conceptual Foundations of Evolutionary Biology
(Chicago: University of Chicago Press, 2004), especially chapter 4, in which they discuss the relationship between statistical formalism and causal analysis. I thank Jim Griesemer for discussions on this point.

40.
William B. Langdon, “Evolution of GP Populations: Price’s Selection and Covariance Theorem,” in
Genetic Programming and Data Structures
, (Norwell, MA: Kluwer Academic Press, 1998) 167–208; M. Van Veelen, “On the Use of the Price Equation,”
Journal of Theoretical Biology
237 (2005), 412–26; T. Day, “Insights from Price’s Equation into Evolutionary Epidemiology,”
DIMACS Series in Discrete Mathematics and Theoretical Computer Science
71 (2006), 23–43; J. W. Fox, “Using the Price Equation to Partition the Effects of Biodiversity Loss on Ecosystem Function,”
Ecology
87 (2006), 2687–96; Stephen C. Stearns, “Are We Stalled Part Way Through a Major Evolutionary Transition from Individual to Group?”
Evolution
61 (2007), 2275–80.

41.
Darwin,
The Origin of Species
, 395.

42.
Robert Trivers,
Social Evolution
(Menlo Park, CA: Benjamin/Cummings, 1985). For a defining volume on what became known as “evolutionary psychology” see Jerome H. Barkow, Leda Cosmides, and John Tooby, eds.,
The Adapted Mind: Evolutionary Psychology and the Generation of Culture
(New York: Oxford University Press, 1992); Sarah F. Brosnan and Frans B. M. de Waal, “Monkeys Reject Unequal Pay,”
Nature
425 (2003), 297–99. For a recent wide treatment of animal morality see Mark Bekoff and Jessica Pierce,
Wild Justice: The Moral Lives of Animals
(Chicago: University of Chicago Press, 2009).

43.
R. Boyd and P. J. Richerson, “Punishment Allows the Evolution of Cooperation (or Anything Else) in Sizable Groups,”
Ethology and Sociobiology
13 (1992), 171–95; E. Fehr and S. Gachter, “Altruistic Punishment in Humans,”
Nature
415 (2002), 137–40; R. Boyd, H. Gintis, S. Bowles, and P. J. Richerson, “The Evolution of Altruistic Punishment,”
Proceedings of the National Academy of Sciences of the United States of America
100 (2003), 3531–35; E. Fehr and U. Fischbacher, “The Nature of Human Altruism,”
Nature
425 (2003), 785–91, are challenged by A. Dreber, D. G. Rand, D. Fudenberg, and M. A. Nowak, “Winners Don’t Punish,”
Nature
452 (2008), 348–51.

44.
See George Loewenstein, Scott Rick, and Jonathan D. Cohen, “Neuroeconomics,”
Annual Review of Psychology
59 (2007), 647–72 for examples. Also, for a window into “positive evolutionary psychology,” see Dacher Keltner’s
Born to Be Good: The Science of a Meaningful Life
(New York: W. W. Norton, 2009), as well as new findings on cooperative infant behavior in Michael Tomassello,
Why We Cooperate
(Cambridge, MA: MIT Press, 2009).

45.
Robert H. Frank,
Passions Without Reason: The Strategic Role of the Emotions
(New York: W. W. Norton, 1988). For an update of this argument from the field, see Frans B. M. de Waal, “Putting the Altruism Back into Altruism: The Evolution of Empathy,”
Annual Review of Psychology
59 (2008), 279–300. For an update from neurophysiology see the work of Joshua D. Green et al., “An fMRI Investigation of Emotional Engagement in Moral Judgment,”
Science
293 (2001), 2105–8, and “The Neural Bases of Cognitive Conflict and Control in Moral Judgment,”
Neuron
44 (2004), 389–400, showing how emotion and moral reasoning relate in the brain. For a recent study on the importance of imitation as an empathy-building mechanism see A. Paukner, S. J. Suomi, E. Visalberghi, and P. F. Ferrari, “Capuchin Monkeys Display Affiliation Toward Humans Who Imitate Them,”
Science
325 (2009), 880–83.

46.
Robert Trivers, “The Elements of a Scientific Theory of Self-Deception,”
Proceedings of the New York Academy of Science
187 (1999), 111–26.

47.
C. Daniel Batson and Laura L. Shaw, “Evidence for Altruism: Toward a Pluralism of Prosocial Motives,”
Psychology Inquiry
2 (1991), 107–22; Lise Wallach and Michael A. Wallach, “Why Altruism, Even Though It Exists, Cannot Be Demonstrated by Social Psychological Experiments,”
Psychological Inquir
y 2 (1991), 153–55; Sober and Wilson,
Unto Others
, presents the case for pluralism.

48.
Charles Darwin,
The Descent of Man, and Selection in Relation to Sex
, selections and commentary by Carl Zimmer (New York: Plume, 2007), 157.

49.
Peter J. Richerson and Robert Boyd,
Not By Genes Alone: How Culture Transformed Human Evolution
(Chicago: University of Chicago Press, 2005).

50.
Trivers did not mean this as a criticism, but rather as a reflection of how others had viewed it. See Hamilton,
Narrow Roads
, 316; for an update of Hamilton’s argument, see Samuel Bowles, “Group Competition, Reproductive Leveling, and the Evolution of Human Altruism,”
Science
314 (2006), 1569. Also J. Heirich, “Cultural Group Selection, Co-evolutionary Processes and Large Scale Cooperation,”
Journal of Economic Behavior and Organization
53 (2004), 3–35.

51.
Claudia Rutte and Michael Taborsky, “General Reciprocity in Rats,”
PloS Biology
5 (2007), 196.

52.
However, see David Sloan Wilson,
Darwin’s Cathedral: Evolution, Religion, and the Nature of Society
(Chicago: University of Chicago Press, 2002), for an argument showing religion functioning as a positive unifying system. From a neuropsychological angle, see also Jonathan Haidt,
The Happiness Hypothesis: Finding the Truth in Ancient Wisdom
(New York: Basic Books, 2005).

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