Wilson replied to Williams and other group selection critics in his own works, including Unto Others: The Evolution and Psychology of Unselfish Behavior and Darwin’s Cathedral: Evolution, Religion, and the Nature of Society.33 In addition to support from Darwin himself, who applied group selection in a limited fashion, Wilson has support from two other evolutionary theorists: Ernst Mayr and Stephen Jay Gould.
Ernst Mayr as Group Selectionist. Ernst Mayr is unquestionably the dean of twentieth-century evolutionary theory. Throughout his long career he has promulgated a limited form of group selection—limited to the human social group. In 1959 he credited J. B. S. Haldane as the first to identify (in 1932) the “population-as-a-whole as a unit of selection” (as group selection was then called), and suggests that the rate of mutation, degree of outbreeding, distance of dispersal, aberrant sex ratios, and other mechanisms favoring either in-breeding or out-breeding are attributes of populations (but not individuals) that would be selected for or against. In 1982 Mayr identified V. C. Wynne-Edwards as the scientist who formalized the group selection process, but still limited it to the deme, or population level, with only a brief mention of social groups. In 1988 Mayr agreed with the critics of group selection when it comes to animal groups, but argued, “there is a great deal of evidence that human cultural groups, as wholes, can serve as the target of selection. Rather severe selection among such cultural groups has been going on throughout hominid history.” In 1997 Mayr made a distinction between soft and hard group selection. Soft group selection “occurs whenever a particular group has more (or less) reproductive success than other groups simply because this success is due entirely to the mean selective value of the individuals of which the group is composed. Since every individual in sexually reproducing species belongs to a reproductive community, it follows that every case of individual selection is also a case of soft group selection, and nothing is gained by preferring the term soft group selection to the clearer traditional term individual selection.” Soft group selection is just individual selection writ large. Hard group selection, by contrast, “occurs when the group as a whole has certain adaptive group characteristics that are not the simple sum of the fitness contributions of the individual members. The selective advantage of such a group is greater than the arithmetic mean of the selective values of the individual members. Such hard group selection occurs only when there is social facilitation among the members of the group or, in the case of the human species, the group has a culture which adds or detracts from the mean fitness value of the members of the cultural group.” This is genuine group selection that differs qualitatively (not just quantitatively, as in soft group selection) from individual organismic selection, and is the subject of so much controversy. 34
Finally, in 2000, in an extensive interview of Mayr for Skeptic magazine, Frank Sulloway and I queried him on group selection and the controversy it has generated. He surprised us when he said: “George Williams and Richard Dawkins have made a mistake, in my opinion, in completely rejecting group selection. But we have to be careful here to define what we mean by a group. There are different kinds of groups. There is one type of group that is a target of selection, and that is the social group. Hominid groups of hunter-gatherers were constantly competing with other hominid groups; some were superior and succeeded and others were not. It becomes quite clear that those groups who had highly cooperative and altruistic individuals were more successful than the ones torn apart by internal strife and egotism.” So the social environment is as important as the physical environment. “The essential point is that if you are altruistic and make your group more successful, you thereby also increase the fitness of the altruistic individual (yourself)!” Critics, we noted, would argue that it is still the individual being selected for these characteristics, not the group. Mayr countered: “There is no question that the groups that were most successful had these individuals that were cooperative and altruistic, and those traits are genetic. But the group itself was the unit that was selected.”35
Stephen Jay Gould as Group Selectionist. Just before his untimely death in May 2002, Stephen Jay Gould witnessed the publication of his magnum opus, The Structure of Evolutionary Theory, in which he defended group selection as a legitimate part of a hierarchical theory of selection that recognizes different levels at which the selection process occurs.36 Gould begins by systematically building upon Darwin’s cathedral, an apt metaphor as his tome begins with an architectural analysis of the Duomo (Cathedral) of Milan, showing how the original fourteenth-century foundational structure was appended over the centuries with spires and pinnacles, such that we can legitimately say a core structure remains intact while the finished building represents a far richer compendium of historical additions. Gould’s mission is not to raze the Darwinian Gothic structure, or to tear down the neo-Darwinian Baroque facades, but to revise, refine, reinforce, and reconstruct those portions of Darwin’s Duomo that have begun to crumble under the weathering effects of a century and a half of scientific research. The foundation of Darwin’s Duomo rests upon three theoretical pillars that form the basis of a hierarchical theory of evolution: agency, efficacy, and scope.
Agency, or the level at which evolutionary change occurs. For Darwin, it is primarily individual organisms that are being selected for or against. Gould proposes a multitiered theory of evolution where change (and selection) occurs on six levels: gene, cell lineage, organism, deme, species, and clade. It is here where Gould defends his own version of group selection—which he calls species selection—without denying the power of Darwinian organismal selection. He does so with two caveats: “First, adjacent levels may interact in the full range of conceivable ways—in synergy, orthogonally, or in opposition. Second, the levels operate non-fractally, with fascinating and distinguishing differences in mode of functioning, and relative importance of components for each level. For example, the different mechanisms by which organisms and species maintain their equally strong individuality dictate that selection should dominate at the organismal level, while selection, drift, and drive should all play important and balanced roles at the species level.”37
Efficacy, or the mechanism of evolutionary change. For Darwin it was natural selection (and its handmaiden, sexual selection, where females, for example, select for or against characteristics that they like or dislike by choosing certain males) that drives organisms to evolve. Gould does not deny the power of natural selection, but wishes to emphasize that in the three-billion-year history of the earth’s rich panoply of life, there is more to the story. On top of the substratum of microevolution (short-term small changes) Gould adds macroevolution—long-term big changes caused by mass extinctions and other large-scale forces of change. To the bottom floor of adaptationism Gould attaches exaptationism—structures subsumed for later uses and whose original adaptive purposes are now lost to history.
Scope, or the range of effects wrought by natural selection. For Darwin, gradual and systematic change extrapolated over geological expanses of time is all that is needed to account for life’s diversity. For Gould, slow and steady sometimes wins the race, but life is also punctuated with catastrophic contingencies that fall in the realm of unique historical narratives rather than predictable natural laws. In Gould’s view, history, not physics, should be evolutionary theory’s model of science.
Revisions to these three branches of agency, efficacy, and scope (while the main Darwinian trunk retains its theoretical power), says Gould, produce a “distinct theoretical architecture, offering renewed pride in Darwin’s vision and in the power of persistent critiques—a reconstitution and an improvement.”38 How can the paradigms of the original Darwinism, the neo-Darwinism of the synthesis, and Gouldian Darwinism coexist peacefully? We can ask the same question with regard to individual versus group selection, or organismal versus species selection as apparently competing paradigms. In science, doesn’t one paradigm displace another in a way that makes them incompatible? No. Paradigms can build upon one anothe
r and cohabit the same scientific niche. Just as the Newtonian paradigm has been reconstituted to include the paradigms of relativity and quantum mechanics, the overarching Darwinian paradigm has been improved by, for example, the subsidiary punctuated equilibrium paradigm, which constitutes an improved reading of the herky-jerky fossil record whose numerous gaps so embarrassed Darwin. (The gaps, say Gould and his cotheorist Niles Eldredge, represent data of a speciation process that happens so rapidly that few “transitional” fossils are left in the historical record.) Darwinian gradualism and individual selectionism can be supplemented with Gouldian punctuationism and group selectionism. How?
Think of species not as billiard balls being knocked about the table of nature willy-nilly, but as polyhedrons, or multifaceted structures (picture an eight-sided die) that sit on a side until nudged by a potent force, and whose internal properties, Gould writes, “‘push back’ against external selection, thereby rendering evolution as a dialectic of inside and outside.” Without discounting the outside, Gould wants us to look again inward (as so many evolutionary theorists did in Darwin’s own day), where the restricting channels of both nature and history direct the selective forces in particular directions. Although individual selection is more potent and common than group selection, the latter may outdo the former under certain conditions, as is the case with the selection for human morality. “Organismic selection may trump species selection in principle when both processes operate at maximal efficiency, but if change associated with speciation operates as ‘the only game in town,’ then a weak force prevails while a potentially stronger force lies dormant. Nuclear bombs certainly make conventional firearms look risible as instruments of war, but if we choose not to employ the nukes, then bullets can be devastatingly effective.”39
Ernst Mayr said we need to be precise in how we define a group. The same is true in defining what we mean by group selection, and how human morality may have evolved—at least in part—as a function of the group selection process. Darwin focused on tribes, and how they differed in terms of patriotism, fidelity, obedience, courage, and sympathy, and how these differences led to the selection of some tribes over others. Mayr talks about groups in terms of their social bonding and interactions, and how this might have led to certain groups being selected for survival over others. Gould’s emphasis on species selection would seem beyond the scope of these more narrowly confined cohorts, but if we think of the rapid extinction of Neanderthals shortly after the arrival of anatomically modern Cro-Magnon humans—considered to be separate species by most paleoanthropologists—this too may be an example of group selection, where the group is the entire species. And the consequences of an entire hominid species disappearing forever off the face of the earth are profound.
APPENDIX II
MORAL AND RELIGIOUS UNIVERSALS AS A SUBSET OF HUMAN UNIVERSALS
The following 202 human universal traits are culled out of anthropologist Donald E. Brown’s original list of 373 human universals, and are directly related to religious and moral behavior. They are presented as a demonstration of the universality of morality, and as further evidence of its evolutionary heritage. For these universals I have added parenthetical notes indicating the relation I believe each has to morality and religion. Brown’s list is in: Donald E. Brown, Human Universals, New York: McGraw-Hill, 1991.
affection expressed and felt (necessary for altruism and cooperation to be reinforced)
age statuses (vital element in social hierarchy, dominance, respect for elder’s wisdom)
anthropomorphization (basis of animism, anthropomorphic gods of Greece/Rome, attribution of human moral traits to the monotheistic God of Jews, Christians, and Muslims)
anticipation (vital for behaviors to have future consequences)
attachment (necessary for bonding, friendship, pro-social behavior)
belief in supernatural/religion (basis of the codification of morality)
beliefs about death (basis for belief in an afterlife)
beliefs about fortune and misfortune (superstition and religion)
binary cognitive distinctions (good and evil, moral and immoral)
biological mother and social mother normally the same person (elemental in inclusive fitness and kin altruism)
childhood fear of strangers (basis of xenophobia)
classification of behavioral propensities (basis of judging moral and immoral behaviors)
classification of inner states (basis of judging moral and immoral traits)
classification of kin (basis of inclusive fitness and kin altruism)
coalitions (foundation of social and group morality)
collective identities (basis of xenophobia, group selection)
conflict (creates moral problems)
conflict, consultation to deal with (resolution of moral problems)
conflict, means of dealing with (resolution of moral problems)
conflict, mediation of (foundation of much of moral behavior)
conjectural reasoning (necessary for moral judgment)
continua (ordering as cognitive pattern)
cooperation (part of altruism)
cooperative labor (part of kin, reciprocal, and blind altruism)
copulation normally conducted in privacy (moral presumption of exclusivity, unique relationship)
corporate (perpetual) statuses (moral ranking of groups)
coyness display (courtship, moral manipulation)
crying (sometimes expression of grief, moral pain)
customary greetings (part of conflict prevention and resolution)
dance (affiliated with many religious ceremonies)
death rituals (awareness of mortality behind many religious beliefs)
decision making (foundation of moral judgment and resolution of moral dilemmas)
decision making, collective (foundation of group moral judgment and resolution of group moral dilemmas)
differential valuations (necessary for moral judgment)
discrepancies between speech, thought, and action (moral intention v. behavior)
distinguishing right and wrong (foundation for all moral judgment and ethical systems)
divination (element in many religious ceremonies)
division of labor by age (form of status and hierarchy)
division of labor by sex (form of status and hierarchy)
dominance/submission (foundation of hierarchical social primate species)
dream interpretation (part of some religious shamanism)
economic inequalities, consciousness of (involved in status and hierarchy disputes)
emotions (necessary for moral sense)
empathy (necessary for moral sense)
entification (treating patterns and relations as things) (makes social moral relations and problems real)
envy (moral trait)
envy, symbolic means of coping with (dealing with moral trait)
ethnocentrism (xenophobia, group selection, war)
etiquette (enhances social relations)
facial communication (necessary for social relations)
facial expression of anger (communication of moral approval/disapproval)
facial expression of contempt (communication of moral approval/disapproval )
facial expression of disgust (communication of moral approval/disapproval)
facial expression of fear (communication of moral approval/disapproval)
facial expression of happiness (communication of moral approval/disapproval)
facial expression of sadness (communication of moral approval/disapproval)
facial expression of surprise (communication of moral approval/disapproval)
facial expressions, masking/modifying of (communication of moral approval/disapproval)
fairness (equity), concept of (foundation of social/moral justice)
family (or household) (the most basic social and moral unit)
father and mother, separate kin terms for (element in awareness of this so
cial/moral unit)
fear of death (foundation of many religious customs and beliefs)
fears (generate much religious and moral behavior)
feasting (part of many religious rituals)
females do more direct childcare (division of labor in social hierarchical species)
figurative speech (symbolic communication necessary for moral reasoning)
folklore (part of gossip)
food sharing (form of cooperation and altruism)
future, attempts to predict (necessary for moral judgment)
generosity admired (reward for cooperative and altruistic behavior)
gestures (signs of recognition of others, conciliatory behavior)
gift giving (reward for cooperative and altruistic behavior)
good and bad distinguished (necessary for moral judgment)
gossip (necessary for monitoring social relations and to assess moral value)
government (social morality)
group living (social morality)
groups that are not based on family (necessary for higher moral reasoning and blind altruism)
healing the sick (or attempting to) (religious ritual)
hope (higher moral reasoning)
hospitality (enhances social relations)
identity, collective (necessary for group moral relations/xenophobia)
imagery (necessary for symbolic moral reasoning and judgment)
incest between mother and son unthinkable or tabooed (obvious evolutionary moral trait)
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