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GROUP SELECTION, SEX, AND FOSSILS

The first use of natural selection to explain the differences among the faunas and floras of different geological times (by means of differential extinction) was in 1832, six years before Darwin discovered its applicability to the origin of species. Charles Lyell (1832, chap. 8-11, e.g., p. 176) was still an anti-evolutionist, for good reasons, but like Darwin he had insights that are still being rediscovered. One of these insights was group selection. This term is applied (Wright, 1945, as "intergroup selection") to selection of populations below, at, and above the species level. It is clearly one of the most important processes of human evolution, although it is usually not recognized (e.g., Thompson, 1972). Different human populations have different rates of increase and decrease for diverse and interacting cultural reasons. Culture is necessarily a group character. This group selection has obviously led to important changes in the composition of the human species, and continues to do so today. Man is probably the best species for the study of group selection within a species. We are not exempt from nature but anesthetized. apomicts are scattered taxonomically, it may be that some revert to sexuality. The second belief above is relevant only in that there is sometimes selective pressure for loss of sexuality. This is proved by the existence of apomicts unless they normally arise by drift. I use sexuality here to exemplify the method clearly and to indicate what sort of results could be found for sexuality with adequate data. Consider a large set of species, some obligatory apomicts and some at least facultatively sexual. The apomicts will have a greater probability of extinction of lineages and the sexual species will have a greater probability of speciation by splitting of lineages. (I temporarily ignore the fact that each individual apomict is a separate lineage.) However, apomicts will sometimes originate from sexual species because of their immediate advantage. We now have the requirements for an equilibrium. In fact that situation is structurally that of the equilibrium between mutation and selection at one locus in haploids (or in diploids without dominance). In the customary symbols, if the frequency of apomicts is low